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Guide to the 


MARINE ISOPOD 
CRUSTACEANS 


of the Caribbean 


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Guide to the 


MARINE ISOPOD 
CRUSTACEANS 


of the Caribbean 


Brian Kensley and 
Marilyn Schotte 


SMITHSONIAN INSTITUTION PRESS 


WASHINGTON, D.C., AND LONDON 


© 1989 by the Smithsonian Institution 
All rights reserved 


Designer: Linda McKnight 
Editor: Nancy Dutro 


Library of Congress Cataloging-in-Publication Data 
Kensley, Brian Frederick. 

Guide to the marine isopod crustaceans of the 

Caribbean / Brian Kensley 

and Marilyn Schotte. 
p. cm. 

Bibliography: p. 

Includes index. 

ISBN 0-87474-724-4 (alk. paper) 

1. Isopoda—Caribbean Sea— 
Classification. 2. Crustacea—Caribbean Sea— 
Classification. I. Schotte, Marilyn. II. Title. 
QL444.M34K434 1989 
595.3'7209153'35—dcl9 88-38647 

CIP 


British Library Cataloging-in-Publication Data 
available 


Manufactured in the United States of America 
10298 SOo 7s 6) Dae 4 mee 
98 97 96 95 94 93 92 91 90 89 


oo The paper used in this publication meets the 
minimum requirements of the American National 
Standard for Performance of Paper for Printed 
Library Materials Z39.48-1984 


Contents 


l Introduction 


] HISTORIC BACKGROUND 
3 GEOGRAPHIC AREA COVERED IN THIS GUIDE 
4. ARRANGEMENT OF THE GUIDE AND HOW TO USE IT 


5 | ACKNOWLEDGMENTS 
Glossary of Technical Terms 
13 Marine Isopods of the Caribbean 


13. ORDER ISOPODA 

15. SUBORDER ANTHURIDEA 

73. SUBORDER ASELLOTA 
107 SUBORDER EPICARIDEA 
114. SUBORDER FLABELLIFERA 
236 SUBORDER GNATHIIDEA 
243 SUBORDER MICROCERBERIDEA 
246 SUBORDER ONISCIDEA 
251 SUBORDER VALVIFERA 


261 Zoogeography 
261 = FAUNAL PROVINCES 
262 ANALYSIS OF THE ISOPOD FAUNA 
266 THE BAHAMAS 


269 BERMUDA 
269 CAVE ISOPODS 


275 Appendix 
277 Literature Cited 
293 Index 


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Introduction 


The title of this work will no doubt raise several questions in many readers’ 
minds: why the Caribbean? why not the Caribbean and the Gulf of Mexico? 
why only the marine isopods? just what is the “Caribbean area”? We hope 
that the answers to some of these (and other) questions will become 
apparent. 

There are several works that already deal with the isopods of the Carib- 
bean, as part of a wider treatment of North American isopods (e.g., 
Richardson, 1905; Schultz, 1969). Why then this “Isopods of the Carib- 
bean”? As partial answer, the following: many new records of isopods from 
the Caribbean region (in its broadest sense) have appeared in scattered pub- 
lications in the last few decades. The time has come to pull these together in a 
single work. The number of marine laboratories in the area has increased, 
with more and more students exploring especially the shallow marine en- 
vironment. A single work on a relatively speciose and abundant group of 
invertebrates would be useful to such investigators, as they build up a com- 
prehensive view of the biology of the region. Concepts of the taxonomy of 
several isopod groups have changed radically over the last few years; again, 
there is obvious utility in having these changes summarized in a single 
source. New species and records are continuously being found. Having a 
single baseline work decreases the time needed for investigating and estab- 
lishing the validity of such records. 


HISTORIC BACKGROUND 


The history of isopod taxonomic research in the Caribbean really starts with 
a worldwide monographic work on the Cirolanidae by Hansen in 1890. In- 
cluded here were about 12 species from the Danish West Indies, now the 
U.S. Virgin Islands. Since then a few major works on Caribbean isopods 
have appeared, such as Moore’s report on the isopods of Puerto Rico (1901) 
and Menzies and Glynn’s report on the same area (1968). Some areas have 
received considerable attention, such as the aforementioned Puerto Rico and, 
more recently, Belize. A list of 116 species of isopods from Cuba (including 
Oniscidea) has been published (Ortiz, Lalana, and Gomez, 1987). At the 
other extreme, there are no records from a number of localities, especially the 


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INTRODUCTION 3 


islands of the southeastern chain of the Lesser Antilles. In total there are 
about 40 publications, varying from descriptions of single species to longer 
works, that deal with isopods from the Caribbean. These publications will be 
encountered in the following guide, under the specific taxa. 


GEOGRAPHIC AREA COVERED IN THIS GUIDE 


The accompanying map (Figure 1) shows the area for which records are 
included in this guide. 

While it may seem logical to include the Gulf of Mexico, and while there 
are several isopod species common to both areas, this has not been done. 
There are relatively few isopod records from the Gulf; undoubtedly a great 
deal of taxonomic work awaits the careful collector in this area. Also, from a 
zoogeographic point of view, separation of the Gulf may be justified. 

Bermuda, on the other hand, situated in the northwestern Atlantic several 
hundred miles off the coast of the United States, is included. This island, 
although remote from the Caribbean, is swept by waters that earlier have 
passed through the Caribbean. Zoogeographically, the shallow-water Ber- 
mudan and Caribbean faunas have much in common. 

While perhaps not strictly in the Caribbean Sea, the Bahamas and the 
Florida Keys are included here, their shallow-water marine faunas being 
overwhelmingly Caribbean in nature. 

Turning to depths limits, within the area under discussion, species from 
the intertidal to 200 meters have been dealt with in some detail. This arbi- 
trary cutoff depth was selected because most Caribbean isopod species in- 
habit relatively shallow depths. About 30 species have been recorded from 
below 200 meters in the Caribbean, many of these known only from the type 
material. A list of species of this very poorly known deeper fauna is included 
here. Without doubt, many species in the deeper waters of the Caribbean 
await discovery. 

A fascinating group of isopods, while not strictly shallow-water marine 
forms, is included. These are the true cave forms, found mainly in the sub- 
orders Anthuridea and Flabellifera. Given the history of the Caribbean from 
the Quaternary to the present, it is not surprising that caves are common 
throughout the region. These caves may be well inland and contain only 
freshwater, but are more commonly anchialine, that is, having some (fre- 
quently subterranean) link to the sea. Less common, and of lesser interest 
from an isopod taxonomist’s point of view, are the fully marine caves in 
direct communication with the sea or, indeed, under the surface of the sea 
itself. 


4 INTRODUCTION 


ARRANGEMENT OF THE GUIDE AND HOW TO USE IT 


A short introduction to the Crustacea Isopoda is provided, followed by a 
glossary of descriptive terms and morphological features used throughout the 
guide (see Figure 2). 

Keys and diagnoses to the suborders and all lower taxa follow. For ease of 
usage, except in the keys, all taxa are presented in alphabetical order, regard- 
less of their phylogenetic relationships. 

Diagnoses are provided for all suborders, families, genera, and species. 
The only exceptions to this are in the suborders Epicaridea and Oniscidea. 
Within each suborder, a key to the families occurring in the Caribbean is 
provided. Similarly, within each family and genus, keys are provided to the 
relevant genera and species, respectively. 

In whatever context, where an author and date appear, a reference to these 
is provided in the Literature Cited section. In some cases, reference is made 
to useful publications such as revisions of families or genera. 

As this is not a textbook on the Isopoda, biological information is generally 
kept to a minimum. In the case of individual species, however, what little 
ecological information is available, is provided. For general texts on biology, 
internal anatomy, physiology, and reproduction, the reader is referred to 
works such as Kaestner (1967), Waterman (1960), Bliss (1982-1985), and 
Schram (1986). 

Within each species discussion, a diagnosis is given, along with maximum 
(total) middorsal lengths for males and females, where known. The diag- 
noses are not exhaustive, but provide only the information needed to distin- 
guish the species. Diagnoses thus vary in length from the statement of a 
single feature to a paragraph concerning several features, depending on the 
understanding and complexity of the taxonomy of the group. In the longer 
diagnoses, morphological features are dealt with in order from anterior to 
posterior on the animal’s body. Records are given, rather than geographical 
distribution, as our knowledge of many species is woefully incomplete. These 
records are given in a roughly north-to-south order; records outside of the 
Caribbean region, as here defined, are given on a separate line. A few species 
not yet recorded from the Caribbean are included, in the strong likelihood 
that they will eventually be found here. The records include depth distribu- 
tion information in meters, where known. Records were taken from pub- 
lished papers; in addition, the collections of the United States National 
Museum of Natural History, Smithsonian Institution, were scoured, and 
many unpublished records from this source are also included. In the “Re- 
marks” section, ecological information such as substrate preferences is given. 
Hosts of parasitic species are given. Formal synonymies are not provided, 


INTRODUCTION 5 


but nomenclatural comments are included in the few cases where a species 
may be known under a more commonly used name. Usually, a figure of the 
entire animal of each species is given. Diagnostic features are usually illus- 
trated. Unless otherwise stated, all illustrations are original and by the au- 
thors, and were made from actual specimens. 

Common and scientific names of fishes that are hosts to parasitic isopods 
are taken from the American Fisheries Society special publication no. 12 
(Robins et al., 1980). 

Finally, a word of warning. Difficulties may be experienced in using the 
keys, for which there may be any of several reasons: characters seen in the 
animal may not clearly conform to those in the key (in which case refer to the 
figures, as well as to good recent descriptions or diagnoses); your material 
may be a new record for the region; or you may have an undescribed species 
(in which case refer to more comprehensive treatments of the group). 


ACKNOWLEDGMENTS 


Much of the material covered in this work comes from the many collectors 
who have deposited specimens from several regions of the Caribbean in the 
collections of the National Museum of Natural History, Smithsonian Institu- 
tion. Rather than risk the unwitting omission of a name, we thank all of these 
individuals collectively. Without their efforts, our knowledge of the Carib- 
bean fauna would be the poorer. 

Material was borrowed from several institutions. We thank the following 
scientists for their assistance in this connection: Jan Stock and Dirk Platvoet 
of the Instituut voor Taxonomische Zoologie, University of Amsterdam; 
Jean Just and Torben Wolff of the Zoological Museum, University of 
Copenhagen; Jacques Forest of the Muséum National d’Histoire Naturelle, 
Paris; Richard Heard of the Gulf Coast Research Laboratory, Ocean 
Springs, Mississippi; Willard Hartman of the Peabody Museum of Natural 
History, Yale University; Herbert W. Levi of the Museum of Comparative 
Zoology, Harvard University; Harold S. Feinberg of the American Museum 
of Natural History, New York; John E. Miller of the Harbor Branch Founda- 
tion, Florida; and Paula M. Mikkelsen of the Indian River Coastal Museum 
at Fort Pierce, Florida. 

Bruce Collette of the National Marine Fisheries Laboratory at the 
Smithsonian Institution assisted with fish names used in this work, for which 
we are grateful. 

We thank the staff of the Scanning Electron Microscope Laboratory of the 
Smithsonian Institution, and especially Susann Braden, who produced the 
electron micrographs used here. 


6 INTRODUCTION 


We are grateful to the Smithsonian Research Opportunities fund, admin- 
istered by David Challinor, then Assistant Secretary for Research, and to the 
Smithsonian’s Caribbean Coral Reef Ecosystems program administered by 
Klaus Riitzler of the Department of Invertebrate Zoology, for the funding of 
several fieldtrips to the Caribbean. The second author acknowledges a finan- 
cial award from the Smithsonian’s Women’s Committee for a grant to facili- 
tate fieldwork. 

Several individuals have provided encouragement, advice, suggestions, 
critical comments, and missed references, all of which have vastly improved 
this work. In this regard we are especially grateful to Thomas E. Bowman, C. 
W. Hart, Jr., Horton H. Hobbs, Jr., and Molly K. Ryan, all of the Depart- 
ment of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution; Dan Adkison, Bass Harbor, Maine (who provided 
considerable assistance with the Epicaridea); Richard Heard of the Gulf 
Coast Research Laboratory, Mississippi; and Paul Delaney of the Los An- 
geles County Museum of Natural History. 

This is Contribution Number 248 of the Caribbean Coral Reef Ecosystems 
(CCRE) program, Smithsonian Institution, supported in part by the Exxon 
Corporation. 


Glossary of ‘Technical ‘Terms 


AESTHETASG. Thin-walled sensory seta usually found on flagellum of 
antennule 

AMBULATORY (as applied to pereopods). Used for walking. 

ANCHIALINE. An aqueous habitat near the sea; referring to saltwater or brackish 
pools fluctuating with the tides, but with no surface connection to the sea. 

ANGULATE. Having an angle or an angular shape. 

ANTENNA. Paired appendage of the third cephalon segment; sometimes referred 
to as antenna 2. 

ANTENNULE. Paired appendage of the second cephalon segment; sometimes 
referred to as antenna 1. 

APICAL. Relating to the apex or tip. 

APPENDAGE. An articulated structure used for feeding, locomotion, sensory 
reception, e.g., mouthparts, antennae, pereopods, pleopods, uropods. 

ARTICLE. A single section of an appendage, with an articulation at one or both 
ends. 

BASIS. Article of appendage adjoining coxa proximally, and carrying endopod 
distally, i.e., article 2 of pereopod. 

BIARTICULATE. Composed of two articles. 

BIDENTATE. Having two teeth. 

BIFID. Divided into two lobes or parts by a cleft. 

BILOBED. Composed of two lobes. 

BIRAMOUS. Composed of two rami or branches. 

BIUNGUICULATE. Having two claws, as in a bifid dactylus. 

CARINA. A keel, or an acute ridge. 

CARINATE. Having one or more carinae or acute ridges. 

CARPOCHELATE. Having a chela or pincerlike structure formed by the seventh 
(dactylus) and fifth (carpus) articles of an appendage. 

CARPUS. Article 5 of pereopod. ; 

CEPHALON. Anterior region of body or head; more correctly the cephalothorax 
in isopods, as the first pereonal segment is usually fused with the head. 

CHELA. Distal pincerlike part of appendage, often formed by a mobile and an 
immobile finger. 

CHELATE. Having a chela; modified to form a pincer. 

CLAVATE. Club shaped; having one end thickened. 

CLYPEUS. Platelike structure of cephalon, anterior to upper lip or labrum, 
sometimes fused with frontal lamina. 

CONGLOBATE. Able to roll up into a ball, as in some sphaeromatid and 
oniscidean isopods. 

CONSPECIFIC. Belonging to the same species. 

CONTIGUOUS. Touching. 


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GLOSSARY 


COPULATORY STYLET. Structure situated on endopod of pleopod 2 in males, 
used for transfer of spermatophore in some species; also referred to as appendix 
masculina. 

CORDATE. Heart shaped in outline. 

coxA. Basal article of an appendage, attached to sternite, sometimes expanded 
into a lateral coxal plate. 

CRENULATE. Having a scalloped edge with rounded teeth, usually used to refer 
to the margin of a structure. 

DACTYLUS. Terminal (7th) article of a pereopod or thoracic appendage. 

DENTATE. Edged with teeth. 

DENTICLE. A small tooth. 

DENTICULATE. Having fine teeth. 

DIGITIFORM.  Fingerlike. 

DISTAL. Situated away from the base or point of origin or attachment. 

EcDysIs. Molting of the integument. 

EMARGINATE. Having the margin concave. 

ENDITE. Medially directed lobe of coxa or basis of an appendage, especially the 
maxilliped. 

ENDOPOD. Inner ramus of a biramous appendage. 

ENTIRE. Complete; usually referring to the margin of a structure that is smooth. 

EPIMERON. Lateral part of a somite. 

EPIPOD. Lateral extension of a protopodite. 

EXCAVATE. Hollowed out. 

EXOPOD. Outer ramus of a paired appendage. 

FALCATE. Sickle shaped; curved and tapering to a point. 

FLAGELLUM. Distal part of antenna or antennule, usually multiarticulate, 
occasionally reduced to one or a few articles. 

FRONTAL LAMINA. Platelike structure of the cephalon immediately anterior to, 
and sometimes fused with, clypeus. 

GENICULATE. Bent at an abrupt angle, as in the body of many arcturid isopods. 

GRANULATE. Having the appearance of bearing beadlike or grainlike 
protuberances; usually applied to the description of a surface. 

HIRSUTE. Bearing hairs (elongate hairs in the case of most isopods). 

HYPOGEAN. Underground. 

IMMERSED. Sunken into, as with one structure into another. 

INCISOR. Cutting process of the mandible, usually dentate, sometimes modified 
for piercing. 

INDURATE. Hardened, usually by calcium carbonate or sclerotized protein. 

INTEGUMENT. Outer covering, e.g., the exoskeleton. 

INTERSTITIAL. Relating to interstices; living in the interstices of sand grains, 
gravel, or rubble. 

ISCHIUM. Article 3 of pereopod. 

LABIUM. Lower lip; usually consisting of a pair of lobes posterior to the mouth. 

LABRUM. Unpaired projection anterior to mouth, attached to the clypeus; upper 
lip. 

LACINIA MOBILIS. Small, usually toothed, process articulating at base of incisor 
in left or both mandibles. 

LAMELLAR. In the shape or structure of a thin plate or lamella. 


9 


10 GLOSSARY 


LAMINA DENTATA.  Serrate platelike structure in the mandible of anthurideans, 
formed by the fusion of spines of the spine-row. 

LANCEOLATE. Lance shaped; narrow and tapering to a point. 

LINGUIFORM. ‘Tongue shaped. 

MANCA. Young of some peracaridean crustaceans (including isopods), lacking 
last thoracic appendage at time of release from broodpouch. 

MANDIBLE. First pair of mouthparts, functioning as jaws, often sclerotized. 

MARSUPIUM. Structure in which eggs are retained by female; the broodpouch. 

MAXILLA (1 and 2). Two sets of paired mouthpart appendages immediately 
posterior to mandible. 

MAXILLIPED. First paired appendage of the thorax; usually incorporated into the 
mouthparts. 

MEDIAN. At, near, or directed toward the middle or midline. 

MERUS. Article 4 of pereopod. 

METAMORPHOSED. ‘Transformed; changed in appearance, structure, or function. 

MESIAL. Near or toward the middle or midline. 

MOLAR. Grinding, and sometimes piercing, structure of the mandible. 

MULTIARTICULATE. Composed of many articles. 

NATATORY. Adapted for swimming. 

OBSOLETE. Becoming vestigial, and losing original function. 

OMMATIDIA. Individual visual components of the compound eye. 

OOSTEGITE. Medially directed lamellar structure arising from coxa of pereopod 
in the female, forming part of the broodpouch or marsupium. 

OPERCULIFORM. In the form of a cover or lid. 

OVATE. Egg shaped or oval. 

PALM. Cutting edge of the propodus, often defined proximally by a spine, in a 
subchelate appendage. 

PALP. Articulated ramus consisting of one to three articles in mandible, of up to 
five articles in the maxilliped. 

PECTINATE. Having teeth like a comb. 

PEDUNCLE. Stalk or proximal part of an appendage, as in antennae. 

PENIAL RAMI. Paired submedian process on sternite 7 of male. 

PEREON. Middle or thoracic region of the body, consisting of seven segments or 
pereonites, first fused with cephalon in isopods. 

PEREONITE. Segment of the pereon. 

PEREOPOD. Paired appendage of the pereon, consisting of seven articles when 
unmodified. 

PILOSE. Covered with short hairs or setae. 

PLEON. Posterior or abdominal region of the body, primitively consisting of six 
segments or pleonites, and bearing paired pleopod and uropod appendages. 

PLEONITE. Segment of the pleon. 

PLEOPOD. Paired appendage of the pleon, five pairs being present in the 
primitive condition. 

PLEOTELSON. Structure resulting from the fusion of the telson and one or more 
pleonal segments. 

PLICATE. Pleated or folded. 

PRANIZA. Juvenile, immature stage of gnathiideans. 

PREHENSILE. Adapted for holding or clinging. 


GLOSSARY 11 


PRODUCED. Extended or lengthened. 

PROPODuUS. Article 6 of pereopod. 

PROTANDROUS. In hermaphroditic forms, becoming a functional male producing 
spermatozoa before becoming a functional female producing eggs. 

PROTOGYNOUS. In hermaphroditic forms, becoming a functional female 
producing eggs before becoming a functional male producing spermatozoa. 

PROTOPODITE. Proximal part of an appendage, consisting of the coxa and basis. 

PROXIMAL. Situated near the point of attachment. 

PYLOPOD. First pereopod of the Gnathiidea, modified to form part of the 
mouthparts. 

RAMUS. Branch of an appendage. 

RENIFORM. Kidney shaped. 

RETICULATE. Resembling or forming a network. 

RETINACULAE. Small hooks on an appendage, used to link the left and right 
members of a pair of appendages. ; 

ROSTRUM. Anterior middorsal projection of cephalon. 

SAGITTATE. Arrow shaped. 

SCLEROTIZED. Hardened, usually with chitin. 

SERRATE. Edged with toothlike projections as in a saw. 

SETIFEROUS. Bearing setae. 

SETOSE. Bearing setae. 

SINUATE. Having a wavy margin. 

sINUOUS. Having curves. 

SOMITE. Body segment, usually having a pair of appendages. 

SPATULATE. Shaped like a spatula. 

SPICATE. Shaped like a spike. 

SPINE-ROW. Row of spines situated between the incisor and molar processes of 
the mandible. 

SPINOSE. Bearing spines. 

STATOCYST. Small saclike sensory organ, often containing granules, used to 
indicate to the animal its orientation. 

STYGOBIONT. Cave organism. 

STYLIFORM. Having a long, slender, stilettolike shape. 

suB-. A prefix indicating “almost” or “just less than,” e.g., submarginal—almost 
on the margin. 

SUBCHELATE. Having a subchela, forming a pincerlike structure, especially by 
the dactylus folding back on the propodus. 

SUTURE. AA line indicating an area of articulation, or of incomplete fusion. 

SYMPOD. Proximal part of an appendage, often formed by the fusion of the coxa 
and basis. 

TELSON. ‘Terminal part of the body, usually bearing the anus. 

THORAX. ‘Tagma or body region between the cephalon and the abdomen. 

TRACHEATE. Bearing tubular respiratory trachea (more correctly pseudotrachea) 
on pleopods, as in Oniscidea. 

TRICUSPID. Bearing three cusps or points. 

TRIDENTATE. Having three teeth. 

TRIFID. Divided into three parts or lobes. 

TRILOBED. Divided into three lobes. 


12 GLOSSARY 


TRISINUATE. Having three curves. 

TRIUNGUICULATE. Bearing three claws, as in a trifid dactylus. 

TRUNCATE. Having the appearance of having been abruptly cut off. 

TUBERCULATE. Bearing knoblike or wartlike prominences or tubercles. 

UNIARTICULATE. Composed of one article. 

UNIRAMOUS. Having one ramus or branch. 

UNIUNGUICULATE. Having a single claw, as in a dactylus. 

UROPOD. Paired pleonal appendage of the last pleonite, usually situated at the 
base of the telson. 


Marine Isopods of the Caribbean 


Phylum Arthropoda 

Superclass Crustacea Pennant, 1777 
Class Malacostraca Latreille, 1806 
Subclass Eumalacostraca Grobben, 1892 
Superorder Peracarida Calman, 1904 
Order Isopoda Latreille, 1817 


DIAGNOSIS Body usually dorsoventrally depressed, occasionally sub- 
cylindrical, rarely bilaterally compressed. Carapace lacking. Antennules and 
antennae uniramous (scale on antenna in some asellotes may represent rudi- 
mentary second ramus). Eyes sessile (although situated on nonmobile stalks 
in some asellotes). Mouthparts consisting of one pair of mandibles, two pairs 
of maxillae, one pair of maxillipeds; latter appendages of first thoracic seg- 
ment fused with cephalon. Mandible usually with palp consisting of one to 
three articles; incisor, lacinia mobilis, and molar usually present; lacinia mo- 
bilis often differing on left and right sides, sometimes absent from right man- 
dible; molar variable. Maxilliped usually consisting of palp of no more than 
five articles, lamellar endite often with coupling hooks, lamellar epipod. Per- 
eonites usually separate, although pereonite 1 sometimes fused with 
cephalon. Coxae of pereopods variously fused with, and forming expanded 
lateral processes of, pereonites. Pereopod | forming additional mouthpart 
(pylopod) only in Gnathiidea. Pereopods generally similar, ambulatory; 
pereopods 1—3 secondarily variously modified and becoming subchelate or 
prehensile; pereopods 4—7 occasionally modified, becoming natatory or pre- 
hensile. Pereopod 7 occasionally not developed (neotenous condition). 
Broodpouch or marsupium formed by varying number of oostegites attached 
ventrally and medially to coxae of pereopods; eggs held in anterior or pos- 
terior pockets or internal pouches in gnathiids and some sphaeromatids. 
Pleon consisting of six pleonites, free or variously fused, plus telson; if one or 
more pleonites fused with telson, resulting structure referred to as pleotelson. 


13 


14 ISOPODA 


Key to suborders of Isopoda 


1. Parasitic on crustaceans; body of 2 nearly always asymmetrical 
PE hed Pere ek, IN eae SIME, Sales hale aiclincd brea Epicaridea 
Free-living or parasitic on fishes; body of ? bilaterally symmetrical, or 
if parasitic: 2 semewhat Gistorted) A505 - pct 40 - = ee epee ne ee 2 


2. Body more or less bilaterally compressed ........... Phreatocoidea* 
Body more or less dorsoventrally depressed or subcylindrical ....... 3 


3. With six pereonites and five pairs of pereopods .......... Gnathiidea 
With seven pereonites and six or seven pairs of pereopods .......... 4 


4. Body usually more than six times longer than wide, subcylindrical, 
uropods never operculiform’ >... 4 e5 Fo <b oe er tocar 5 
Body usually less than six times longer than wide, usually 
dorsoventrally depressed; if subcylindrical, uropods operculiform .. 6 


5. Uropodal exopod often folding dorsally over pleotelson; rarely 
imterstitial farmisys). eset s orden ake 3s AS bicenneted oe beve Anthuridea @!¢ 
Uropods terminal, exopod lacking; minute interstitial forms 
cealidinttiansd sie Baas viasteetide ahyetien see syeleieg Microcerberidea 


6. Antennules minute; terrestrial forms, with pleopods tracheate 


SN pe Ce ar reer ROR Al SST ard ry Sowa Ue ros pie Bait ioe3 Oniscidea 
Antennules rarely minute; aquatic forms, pleopods never tracheate .. 7 
: : : 75"\ 
7. Uropods ventral, operculiform, covering pleopods .......... Valvifera ?'~ 
Uropods never operculitorm;over pleopods) 0 .)5.00. 21st ce eee 8 


8. Uropods lateral or ventrolateral, forming tailfan with pleotelson; 


pleopods 1 and 2 rarely operculiform <......... 0. 6c. on Flabellifera ¢ !\\ 
Uropods terminal or subterminal; pleopods | and 2 variously 
aperculilorian specu sna gttie asses eso ce ei eee CeCe eae Asellota 9+ ’ 


* The suborder Phreatocoidea contains freshwater forms, and has a 
Gondwanian distribution, primarily in the Southern Hemisphere. 


Pleopods (on pleonites 1—5) biramous, lamellar, primarily for respiration; 
anterior pleopods occasionally operculiform. Pleopod 2 in male (and occa- 


ANTHURIDEA 15 


sionally also pleopod 1 in Oniscidea and Asellota) with endopod bearing 
copulatory stylet. One pair of uropods on pleonite 6. Young leave brood- 
pouch as manca, i.e., resembling adult but lacking pereopod 7; in Epi- 
caridea, manca stage represented by epicaridium stage; latter transforms 
into microniscium and then cryptoniscium stage, before becoming adult. 


Suborder Anthuridea Leach, 1814 


DIAGNOSIS Body generally elongate and subcylindrical. Eyes absent in 
some genera. Antennular peduncle of three articles; antennal peduncle of five 
articles. Mandible with palp of one to three articles, or absent; body of mand- 
ible either styliform and lacking molar and lacinia mobilis, or with molar 
variously specialized or reduced, lacinia mobilis absent, and spine-row modi- 
fied to form platelike lamina dentata. Maxilla 1 with inner ramus reduced, 
outer ramus slender. Maxilla 2 rudimentary. Maxilliped variable, with palp 
of one to five articles, endite present, modified, reduced, or absent. Pereonite 
1 free. Pereopod 1, or pereopods 1—3 subchelate; pereopods 4—7 generally 
ambulatory. Pleonites 1—5 free or fused, pleonite 6 partly or completely fused 
with telson. Pleopods 1—5 similar, or pleopod 1 variously modified to form 
operculum. Uropodal exopod often folded dorsally over pleotelson. 
Pleotelson with pair of statocysts, with single statocyst, or lacking statocysts. 


REMARKS Protogyny has been demonstrated in several species of An- 
thuridea. The order of development in these cases is: egg, manca (both in the 
broodpouch), immature subadult, ovigerous female, premale, male, with 
varying numbers of molts between each stage. At least one molt takes place 
between ovigerous female and premale, the latter being distinguished by the 
loss of the oostegites and by the elongation of, and acquisition of more flagel- 
lar articles in, the antennule. One or two molts take place between premale 
and sexually mature male, the latter being characterized by the possession of 
elongate antennular flagella bearing dense whorls of aesthetascs, a more set- 
ose and/or spinose pereopod 1, and sometimes by an elongation of the pleon 
and uropods. In some genera, the males have somewhat atrophied 
mouthparts, suggesting that they do not feed at this stage. As a result of this 
seemingly widespread protogyny, sex ratios are strongly biased toward 
females, and in several species males are not yet known. 

The number of families in the suborder Anthuridea has not been settled. 
At present, three families are recognized. Doubtless, further families will be 
defined and the genera reshuffled. 


16 ANTHURIDEA * ANTHURIDAE 


Key to families of Anthuridea 


1. Mouthparts adapted for piercing and sucking, together forming 
conclike-structire! ..29 744 sv i beeetere as. Paranthuridae p 64 
Mouthparts adapted for cutting, lamina dentata and molar usually 


presemty:. 0. casktenes Hace ere: cnet te cn salsa ote 2 
2. Pereopod | subchelate, with propodus expanded; pleonites generally 
fused; if free; much: shorter than wide ©. .... ..\0.2 284 20a Anthuridae J 
Pereopods 1-3 subchelate, subsimilar; pleonites free, often as long as 
WIE IA EL TAIL ae rte coe ens SRE Grate ata eae a Oaee ror Hyssuridae pco 


Family Anthuridae Leach, 1814 


DIAGNOSIS Mouthparts adapted for cutting. Pereopod | usually markedly 
different from remaining pereopods, subchelate with propodus more or less 
inflated. Exopod of pleopod | operculiform, covering remaining pleopods. 
Pleonites 1—5 fused, with fusion marked ventrolaterally by short slits, occa- 
‘ sionally with dorsal grooves marking lines of fusion, or free; if free, length of 
each pleonite much less than width. Pleotelson with pair of statocysts, or 
single medial statocyst, or lacking statocysts. 


Key to genera of Anthuridae 


1. Pleopod 1, both rami contributing to operculum ................. 2 
Pleoped I, only exopod operculiform, 2... 2/5. 605+ vata -s---%-=5 6 

2. Antennal peduncle bearing serrate process ............-- Licranthura +3 
Antennal peduncle lacking any serrate process ..............-.+-- 3 

3.. Pleopod Jim: Y rami to Some depree fused 2... 36. Eisothistos 32 
Pleopod: iin; 2; ram. trees") - 5.04 seb om se cee ee eee 4 


4. Pereopods | and 2 subchelate, of similar size, propodi not noticeably 
inflated (12 24! < ooeigads Saba 5 aglaw Sete ake eee) «Site 5 
Pereopod | much larger and propodus more expanded, than 
pereopod 2 s2...ssigekite wadeanst anh eee ernie ee Minyanthura 


Amakusanthura 17 


5. Integument noticeably pitted; mandible on one side lacking molar, on 


Other side wath spleate Molar TH. Re en ek Apanthuroides  p- ab 
Integument not pitted; mandibles similar on both sides Chalixanthura_ p- 2F 
Be Melson awathissiale/SEALGEVSE (oe ep a am ay bine oe wt ¥en spe ens Anthomuda 
Telson with two statocysts, or lacking statocysts ................. i 
7+ Pereopods 4—7; carpi roughly rectangular 2... .:. ..... ..)R¥ekas Jed 8 
Pereopods 4—7, carpi triangular, with anterior margin considerably 
Shorter than: pOstenior Margihy 205 acd ase ariewid des 1 Wasorw were 9 
3: Maxallipedal palpof tworarticless: “220 ys Pali. css Haliophasma 
Masallipedal palp*of three articles 2... 222 2.0.05%5 sic. s. Malacanthura 
9. Maxillipedal palp of one article; mandibular palp of one article 
BR pe RNC cle S aeRO PANEL dears GIES efeal kay Gee cts Gores aia SP Pendanthurag. sc 
Maxillipedal palp of more than one article; mandibular palp of more 
HINARI CONG ATERELE? (os oh osc sb NEI te ela capn Mes Mp2 ear a tered a Ry ee aerate 10 
LO: ¢Miasallipedal palpiof two articles \...4.: 2 sls ds25 oh see oe dete ee oe 1] 
Maxllipedalspalprot three articles... 2.2442 <slc ss cane dees oe « 12 
iesMandibularpalp of two,articles,. .. 2... 220 Mataania hones Cortezura P- 24 
Mancibular palprot threearucles: . (v6). 2b. ana son ois sh «as Cyathura f. 3] 
12. Cephalon with midventral process at base of mouthparts .. Skuphonura 
Cephalon lacking midventral process')!.26°.4.0:0).g0ditcue half 6 i 
13. Pleon lacking dorsal grooves or lines indicating boundaries of fused 
pleonites; species-specific persistent dorsal pigment pattern usually 
PUES IU eect hs erent Heh hers iectrocte ect the arts evap any cdl osene -adbvatone Mesanthura 
Pleon with dorsal grooves or lines indicating boundaries of fused 
pleonites; persistent pigment pattern lacking ................. 14 
14. Pleon with complete dorsal lines separating pleonites 
Da oie cane eects eta Sita cc Nar eT ae slee oe edi ee, SaaS Apanthura po 
Pleon with incomplete dorsal line between pleonites 4 and 5 
PP Se eee EN tN Gt aE bette ocak epee ae ate Amakusanthura ,\*°'* 


Amakusanthura Nunomura, 1977 


DIAGNOSIS Integument sometimes with pigment. Eyes present. Antennular 
flagellum of three articles. Antennal flagellum of two to four articles. Man- 
dibular palp of three articles, terminal articles bearing distal spines. Max- 


18 ANTHURIDEA * ANTHURIDAE 


illipedal palp of three articles; endite small or absent. Pereopod 1, propodal 
palm with step or tubercle. Pereopods 4—7, carpus triangular. Pleonites 
short, pleonites 1—4 fused, boundaries demarked by complete dorsal folds, 
pleonites 4 and 5 separated only by lateral fold, not demarked dorsally; 
pleonite 6 dorsally demarked from telson. Pleopod | exopod operculiform. 
Uropodal exopod often notched or excavate distally. Pleotelson with two 
basal statocysts. 


Key to species of Amakusanthura 


1. Telson thickened, with raised area at midlength, widening and sloping 
posteroventrally; uropodal exopod distally shallowly excavate 
REE Ae Mere metre eT Sk FAO HUM VE End ie CEN magnifica 
Telson dorsally flat, not thickened; uropodal exopod distinctly notched 
on barely excavate distallys pjis sss ots oe ao one my ere te 2 


2. Integument pigmented; uropodal endopod length 1.5 times basal width 


sinat idl dfn Reka Betas ee CSU Gs ENE VA SEUSS a SE ace tL MSIE RMN Rear Re eaae a signata 

Imtesument not pigmented)... inside sedlk eip atl eieh etd ree 3 

3: Uropodal) exopod! distimetly notched... sic oc \in. - eae et Oe. eee 4 
Uropodal exopod, outer margin weakly excavate; mandibular palp 

anticle:J*bearing three spines mess aaselt cielo icin) eect gee lathridia 

4. Mandibular palp article 3 bearing three spines .............. significa 

Mandibular palp article 3 bearing two spines ............-. geminsula 


Amakusanthura geminsula (Kensley, 1982) 
Figure 3A—E 


DIAGNOSIS. 2: 8.1 mm. Shallow middorsal pit on pereonites 4—6, appearing 
chalky white in life. Antennular flagellum of three articles. Antennal 
flagellum of two articles. Mandibular palp article 3 bearing two spines. Max- 
illiped with short slender endite; terminal palp article set obliquely at outer 
distal angle of penultimate article. Pereopod 1, propodal palm with step; 
carpus distally a rounded lobe. Uropodal exopod with notch. Pleotelson 
tapering in posterior half to subtruncate apex. d: 4.8 mm. Eyes larger than 
?. Antennular flagellum of 8—9 articles. Pereopod 1, carpus with distal lobe 


Amakusanthura geminsula 19 


Figure 3. Amakusanthura geminsula: A, 2; B, pereopod 1; C, telson; D, uropodal 
sympod and endopod; £, uropodal exopod. Amakusanthura lathridia: F, pereopod 1; 
G, telson; H, uropodal exopod; J, uropodal sympod and endopod. 


narrowly rounded; propodal palm with rounded tubercle; irregular band of 


setae on mesial surface. 


RECORDS Carrie Bow Cay and Twin Cays, Belize, intertidal to 1.5 m; 
Jamaica. 


20 ANTHURIDEA * ANTHURIDAE 


Amakusanthura lathridia (Wagele, 1982) 
Figure 3F—I 


DIAGNOSIS @ 2.7 mm. Antennular flagellum of three articles. Antennal 
flagellum of three articles. Mandibular palp article 3 bearing three spines. 
Maxilliped lacking endite. Pereopod 1, propodal palm with distal step; 
carpus with triangular posterodistal lobe. Uropodal exopod apically acute, 
outer margin slightly excavate, sinuate; endopod length slightly less than 
twice basal width. Pleotelson elliptical, widest at about midlength, margins 
weakly serrate. 


RECORDS Cuba, interstitial beach sand just above water line. 


Amakusanthura magnifica (Menzies and Frankenberg, 1966) 
Figures 4, 5 


DIAGNOSIS 2: 13.9 mm. Maxilliped with short endite, terminal article of 
palp short, set obliquely at outer distal angle of penultimate article. Pereopod 
1, propodal palm with strong tubercle; carpus with low distal sclerotized 
lobe. Uropodal exopod ovate, with shallow distal notch; endopod twice 
longer than wide. Pleotelson with posterior margin evenly rounded to sub- 
truncate; dorsally with raised area at midlength, broadening and sloping 


Figure 4. Amakusanthura magnifica: A, 
pereopod 1; B, telson; C, uropodal sympod 
and endopod (setae omitted); D, uropodal 


exopod (setae omitted) 


Amakusanthura signata 21 


Figure 5. Amakusanthura magnifica: A, 
pleonites in lateral view; B, pleon in dorsal 


view. 


away posteriorly. d: 10.3 mm. Antennular flagellum of 24 articles. Pereopod 
1, carpus with sclerotized distal rounded lobe; propodal palm with strong 
sclerotized tubercle longer than in 2; numerous setae on mesial surface. 
Pleopod 2, copulatory stylet of endopod not reaching beyond ramus. 


RECORDS Off Georgia, 17-137 m; off Florida, 7-11 m; Cuba; Gulf of 
Mexico. 


Amakusanthura signata (Menzies and Glynn, 1968) 
Figure 6A—E 


DIAGNOSIS @: 4.9 mm. Integument with strong patches of pigment on 
cephalon; pigment sparse on pereonites; two bars on pleonite 6. Antennular 


22 ANTHURIDEA * ANTHURIDAE 


Figure 6. Amakusanthura signata: A, 2; B, telson; C, pereopod 1; D, uropodal 
sympod and endopod; £, uropodal exopod. Amakusanthura significa: F, telson; G, 
uropodal sympod and endopod; H, uropodal exopod; /, pereopod 1. 


flagellum of three articles. Antennal flagellum of three articles. Mandibular 
palp article 3 bearing three spines. Maxilliped with short endite not reaching 
base of palp article 2. Pereopod 1, carpus with strong subacute sclerotized 
lobe distally; propodal palm with transparent convex flange and step. 


Anthomuda stenotelson 23 


Uropodal exopod with strong distal notch; endopod length about 1.5 times 
basal width. Pleotelson widest in posterior half, tapering posteriorly to shal- 
lowly notched apex. ¢: 4.5 mm. Eyes larger than in ¢. Antennular flagellum 
of nine articles. Pereopod 1, carpus with strong distal lobe, propodal palm 
with step, convex transparent flange, and numerous setae on mesial surface. 
Pleopod 2, copulatory stylet of endopod reaching well beyond ramus. 


RECORDS Cuba; Puerto Rico, intertidal to 1.5 m; Carrie Bow Cay, Belize, 
intertidal to 24 m. 


Amakusanthura significa (Paul and Menzies, 1971) 
Figure 6F-I 


DIAGNOSIS @ 5.0 mm. Antennular flagellum of three articles. Antennal 
flagellum of four or five articles. Mandibular palp article 3 bearing three 
spines. Maxilliped lacking endite. Pereopod 1, propodal palm with step; 
carpus distally rounded. Uropodal exopod ovate, with distal notch; endopod 
length slightly more than twice basal width. Telson elliptical, posterior mar- 
gin narrowly rounded. 


RECORDS Off Venezuela, 95 m. 


Anthomuda Schultz, 1979 


DIAGNOSIS. Eyes present. Mandibular palp of three articles. Maxillipedal 
palp of four articles; endite reaching well beyond base of palp article 2. Per- 
eopods | and 2 similar, subchelate. Pleonites 1—6 short, free; pleonite 6 dor- 
sally demarked. Telson with single semiovate hollow representing statocyst. 


Anthomuda stenotelson Schultz, 1979 
Figure 7A 


DIAGNOSIS @ 8.8 mm. Antennular flagellum of three or four articles. An- 
tennal flagellum of six articles. Pereopods | and 2 similar, propodi somewhat 
inflated; pereopod 1, propodal palm straight, unarmed; pereopod 2 propodal 
palm with three sensory spines. Pereopods 4—7 with rectangular carpi. 
Pleonite 6 free, posterior margin broadly bilobed. Uropodal exopod elongate- 
elliptical, distally narrowly rounded; endopod length 2.5 times basal width, 
distally broadly rounded. Telson narrowly lanceolate, posteriorly narrowly 
rounded, with single open hollow statocyst. 


RECORDS Off Bermuda, 90 m. 


Figure 7. Anthomuda stenotelson: A, 2°. Apanthura cracenta: B, 2; C, telson; D, 
uropodal endopod and sympod; £, uropodal exopod. Apanthura crucis: F, uropodal 
exopod; G, uropodal endopod and sympod. Apanthura harringtoniensis (from Wagele, 
1981): H, uropodal endopod and sympod; /, uropodal exopod. 


Apanthura cracenta 25 


Apanthura Stebbing, 1900 


DIAGNOSIS Integument sometimes pigmented. Eyes present. 2: Antennu- 
lar flagellum of three articles. Antennal flagellum of two to four articles. 
Mandibular palp of three articles, terminal article bearing distal spines. 
Maxillipedal palp of three articles; endite small, or lacking. Pereopod 1, pro- 
podal palm usually with step or tubercle; propodus inflated. Pereopods 4—7, 
carpus triangular. Pleonites 1—5 fused; pleonite 6 dorsally demarked. 
Pleopod 1, exopod operculiform. Uropodal exopod ovate, sometimes distally 
notched or excavate. Pleotelson with two basal statocysts. ¢ antennular 
flagellum of about 10 articles. 


REMARKS Differentiation of species, especially if long preserved and the 
pigmentation is lost, depends on subtle features of the mandible, maxilliped, 
pereopods, and pleotelson. 


Key to species of Apanthura 


i, Uropodal'exopod distally notched or‘excavate ...... 2. feos ee. ees 2 
Uropodal exopod distally entire; uropodal endopod length subequal to 
WIE D oa Cie ee Fe Ae gE Se ee Ae ieda Mintel act crucis 


2. Uropodal exopod distally notched; uropodal endopod length 2.4 times 
ate tuyaniel igs os Ns Sess Spal Secgced SS atecun agai Sepe raee, iS sBe a ntte cracenta 

Uropodal exopod faintly excavate or sinuate; uropodal endopod length 
abouimiavmice Hasaliwad the sa.1.21 aca 2. Sc cacucray Pods sigra harringtoniensis 


Apanthura cracenta Kensley, 1984 
Figure 7B—E 


DIAGNOSIS @: 4.6 mm. Antennular flagellum of three articles. Antennal 
flagellum of two articles. Mandibular palp article 3 bearing four spines. 
Maxilliped with short rounded endite; terminal palp article set at oblique 
angle on, and less than half length of penultimate article. Pereopod 1, carpus 
triangular with acute sclerotized tip overlapping base of palm; propodal 
palm with rounded tubercle near midlength. Uropodal exopod deeply 
notched; endopod ovate, length 2!/3 basal length. Pleotelson lanceolate. d: 
3.8 mm. Antennular flagellum of six articles. Antennal flagellum of two arti- 
cles. Pleopod 2, copulatory stylet of endopod reaching by half its length be- 
yond ramus. 


26 ANTHURIDEA * ANTHURIDAE 


RECORDS ‘Turks and Caicos Islands, 1 m; Carrie Bow Cay, Belize, interti- 
dal on reef crest, 2 m. 


Apanthura crucis (Barnard, 1925) 
Figure 7F,G 


DIAGNOSIS ?: 5.9 mm. Antennular flagellum of four articles. Antennal 
flagellum of four articles. Mandibular palp article 3 with eight spines. Max- 
illiped lacking endite; terminal palp article more than half length of penulti- 
mate article. Pereopod 1, propodal palm with low rounded tubercle at mid- 
length. Uropodal exopod ovate, outer margin evenly convex; endopod 
subcircular. Pleotelson anteriorly narrow, widest at midlength, posterior 
margin broadly rounded. ¢: 6.2 mm. Antennular flagellum of 12 or 13 arti- 
cles. Antennal flagellum of four articles. Pereopod 1, propodal palm with low 
rounded tubercle; dense band of setae on mesial surface. Copulatory stylet on 
endopod of pleopod 2 reaching a third of its length beyond ramus. 


RECORDS ‘Turks and Caicos Islands, 1 m; St. Croix, U.S. Virgin Islands, 8 
m. 


Apanthura harringtoniensis Wagele, 1981 
Figure 7H,I 


DIAGNOSIS @ 6.0 mm. Antennular flagellum of four articles. Antennal 
flagellum of four articles. Mandibular palp article 3 bearing two spines. 
Maxillipedal endite short, not reaching base of palp article 2. Pereopod 1, 
propodal palm stepped; carpus distally bluntly triangular. Uropodal exopod 
apically acute, outer margin distally sinuate; endopod length about twice 
basal width. Pleotelson with posterolateral margins faintly denticulate; apex 
subtruncate and bearing several setae. 


RECORDS Harrington Sound, Bermuda. 


Apanthuroides Menzies and Glynn, 1968 


DIAGNOSIS’. Eyes present. Mandibular palp of three articles; body of mand- 
ible anteriorly produced, molar absent on left side, present as narrow 
spikelike process on right. Maxillipedal palp of three articles; endite present, 
reaching palp article 2. Pereopod 1, propodus barely expanded, palm un- 


Chalixanthura Qi, 


armed. Pereonite 7 less than half length of pereonite 6. Pereopods 4—7, carpi 
with free anterior margin shorter than posterior margin. Pleonites 1—5 fused, 
6 fused with telson. Pleopod 1, both rami forming operculum. 


Apanthuroides millae Menzies and Glynn, 1968 
Figure 8 


DIAGNOSIS @: 4.5 mm. Integument with diffuse brown pigmentation and 
numerous shallow pits. Antennular flagellum of four articles. Antennal 
flagellum of seven articles. Pleotelson with strong middorsal ridge. d: 5.0 
mm. Integumental pigment stronger than in @, in irregular brown patches. 
Eyes larger than in 2. Antennular flagellum of six or seven articles. Antennal 
flagellum of seven articles. Pereopod 1, propodal palm armed with five 
fringed spines. 


RECORDS Carrie Bow Cay, intertidal to 30 m; Puerto Rico, intertidal. 


REMARKS The highly modified mandible probably indicates some spe- 
cialized form of feeding, but this has yet to be discovered. 


Chalixanthura Kensley, 1984 


DIAGNOSIS Eyes present, enormously enlarged, especially ventrally in 6, 
resulting in mouthpart reduction. Mandibular palp of three articles; molar 
small, lamina dentata, and incisor present. Mouthparts reduced in 6. Per- 
eopods 1-3 similar, propodi barely inflated. Pereopods 4—7, carpi triangular. 
Pleopod 1, exopod broader than endopod, operculiform, or both rami form- 
ing operculum. Pleopods 2—5, exopods biarticulate. Pleonites short, free, 
longer in d than in 2. Uropodal rami, margins moderately to strongly in- 
cised or serrate. Telsonic margin serrate posteriorly; statocysts lacking. 


Key to species of Chalixanthura 


1. Body pigmented; uropodal exopod deeply incised ........... scopulosa 
Body not pigmented; uropodal exopod weakly serrate .......... lewisi 


Figure 8. Apanthuroides millae: A, 2; B, right mandible; C, left mandible; D, 
maxilliped; £, uropodal exopod; F, pleopod 1; G, pereopod 1. 


Cortezura 29 


Chalixanthura lewisi Kensley and Snelgrove, 1987 
Figure 9A—D 


DIAGNOSIS @: 3.1 mm. Integument lacking pigment. Antennular flagellum 
of three articles. Antennal flagellum of seven articles. Maxillipedal endite 
large, apically acute. Pereopod | propodus slightly expanded, palm with few 
(3) simple setae. Pereopod 7 propodus with two elongate anterodistal fringed 
spines. Uropodal and telsonic margins serrate. d: 2.2 mm. Antennular 
flagellum of 11 articles. Antennal flagellum of six articles. Maxilliped re- 
duced, lacking endite. Pereopod 1, propodal palm with two spines plus seven 
spines on mesial surface. Pereopod 7 as in @. 


RECORDS Barbados, in Madracis mirabilis coral, 9-15 m. 


Chalixanthura scopulosa Kensley, 1984 
Figure 9E—J 


DIAGNOSIS ¢: 2.5 mm. Antennular flagellum of three articles. Antennal 
flagellum of seven articles. Maxillipedal endite short, not reaching base of 
palp article 4. Pereopod 1, propodal palm unarmed. Pereopod 7, propodus 
with single elongate anterodistal fringed spine. Uropodal exopod margin de- 
eply incised; endopod ovate, margins serrate. Telson elongate-ovate, pos- 
terior margin serrate. d: 2.6 mm. Eyes considerably larger and with more 
ommatidia than in 2. Antennular flagellum of seven articles. Antennal 
flagellum of four articles. Pereopod 1 propodal palm with three sensory 
spines. Uropodal exopod deeply incised. 


RECORDS Carrie Bow Cay, Belize, 0.1 m. 


Cortezura Schultz, 1977 


DIAGNOSIS Eyes small, weakly pigmented. Antennular flagellum of two 
short articles. Antennal flagellum of single article. Mandibular palp of two 
articles. Maxillipedal palp of two articles, terminal article small; short endite 
present. Pereopod 1, propodus inflated. Pereopods 4—7 with carpus having 
anterior margin shorter than posterior. Pleonites 1—5 fused; pleonite 6 dor- 
sally demarked. Pleopod 1, exopod operculiform. Pleotelson with two basal 
statocysts. 


Figure 9. Chalixanthura lewisi: A, 36; B, 2; C, uropodal exopod; D, pleopod 1. 
Chalixanthura scopulosa: E, 3; F, 2; G, maxilliped; H, pereopod 1; J, pleopod 1; J, 
uropodal exopod. 


Cyathura (Cyathura) ai 


Cortezura confixa (Kensley, 1978) 
Figure 10 


DIAGNOSIS 2 13.4 mm. Antennular peduncle articles 1 and 2 each with 
clump of ventrally directed setae. Antennal peduncle article 2 and antennu- 
lar peduncle article 1 locked together. Mandibular palp of two articles, distal 
article twice length of proximal. Pereopod 1, propodus expanded, especially 
posteriorly; palm concave, with strong tubercle and irregular band of setae 
on mesial surface. Uropodal exopod ovate, apically rounded; endopod length 
1.5 times greatest width. Telson ovate, posteriorly faintly narrowed, prox- 
imally thickened, with faint ridges diverging posteriorly. 


RECORDS Cubagua Island, Venezuela, 4-10 m. 


REMARKS This species was described under the generic name Venezanthura 
Kensley (1978). The only other known species of Cortezura is the type of the 
genus, C. penascoensis Schultz, 1977, from California. 


Cyathura Norman and Stebbing, 1886 


DIAGNOSIS. Eyes present or absent. Antennular flagellum of one to three 
articles in 2. Antennal flagellum of one to three articles. Mandibular palp of 
three articles. Maxillipedal palp of two articles; endite reduced or absent. 
Pereopod 1, propodus inflated. Pereopods 4—7, carpi triangular. Pleopod 1 
exopod operculiform. Pleonites 1—5 short, fused; pleonite 6 fused with telson, 
sometimes dorsally demarked. Pleotelson with two basal statocysts. 


Key to subgenera of Cyathura 


1. Pleonite 6 not dorsally demarked from telson; articulation of uropodal 
exopod very short; exopod not adpressed to telson dorsally 
SES Sts Dade A Le BR oe Seth. wen Peis eae ieee Stygocyathura 
Pleonite 6 dorsally demarked from telson; articulation of uropodal 
exopod relatively elongate; exopod adpressed dorsally to telson 
ies ebb te mntin see NN aie tes ANE alata oe Sate eral, HARMS Cyathura 


Cyathura (Cyathura) Norman and Stebbing, 1886 


DIAGNOSIS Integument usually strongly pilose or setose; eye and body pig- 


Figure 10. Cortezura confixa: A, 9; B, cephalon and pereonite 1, lateral view 
(pereopod | removed); C, antennule; D, pleon, lateral view; £, mandible; F, 


maxilliped; G, pereopod 1; H, pereopod 7. 


Cyathura (Stygocyathura) 33 


mentation usually present. Pereopod 1, propodal palm armed with tubercle. 
Pleopod 1, protopod with retinaculae. Uropodal exopod articulation relative 
elongate along lateral margin of sympod; exopod well developed, ovate. 
Pleonite 6 dorsally demarked from telson. Marine or estuarine forms. 


Cyathura (Cyathura) cubana Negoescu, 1979 
Figure 11A,B 


DIAGNOSIS Ovigerous 2: 7.0 mm. Antennular flagellum of two articles. An- 
tennal flagellum of one article. Pereopod 1, propodal palm with rounded lobe 
in proximal half. Maxillipedal palp with distal article 0.34 times length of 
proximal article; small rounded endite present. Dorsal pigmentation consist- 
ing of irregular brown mottling. d: 5.5 mm. Antennular flagellum of four 
articles. Antennal flagellum of three articles. Pereopod 1, propodal palm with 
rounded lobe in proximal half. Copulatory stylet elongate-cylindrical, api- 
cally narrowed and flexed. 


RECORDS Cuba, in mangroves, 2.5—7.0 m; Salt Creek, Belize, in man- 
groves, 1.5 m. 


Cyathura (Stygocyathura) Botosaneanu and Stock, 1982 


DIAGNOSIS. Eye and body pigmentation absent. Body sparsely pilose or set- 
ose. Tendency toward elongation of some appendages, especially propodus of 
pereopods 2—7. Pereopod 1, propodal palm lacking strong tubercle. Pleopod 
1, protopod lacking retinaculae. Pleonite 6 fused with telson, not dorsally 
demarked. Uropodal exopod with very short articulation on sympod, not 
adpressed dorsally to telson. Cave or hypogean forms. 


REMARKS _ The ten species of Stygocyathura from the area covered in this work 
are morphologically very similar, with specific differences, although real, 
being very subtle. A dichotomous key would be cumbersome and require 
considerable dissection of mouthparts. The copulatory stylet of the male pro- 
vides a valuable specific feature but males are not always available. Instead 
of a key, we have provided a list of species with their total lengths and lo- 
calities (Table 1). Given the very restricted distribution of these cave species, 
material from localities not listed here should be treated as potentially un- 
described, and the material compared with descriptions, especially those of 
Botosaneanu and Stock (1982). 


34 ANTHURIDEA * ANTHURIDAE 


J 


Figure 11. Cyathura (Cyathura) cubana: A, 2; B, pereopod 1, 2. Cyathura 
(Stygocyathura) cuborientalis: C, pereopod 1, 2; D, telson. Cyathura (Stygocyathura) 
curassavica: EF, pereopod 1, 2; F, telson. Cyathura (Stygocyathura) hummelincki: G, 


pereopod 1, @; H, telson. Cyathura (Stygocyathura) motasi: I, telson; J, pereopod 1, 
ae 


Cyathura (Stygocyathura) cuborientalis Botosaneanu and Stock, 1982 
Figure 11C,D 


DIAGNOSIS 2 6.8 mm. Pereopod 1, propodal palm straight, bearing about 


Cyathura (Stygocyathura) hummelincki 215) 


TABLE |. CARIBBEAN SPECIES OF Cyathura (Stygocyathura), THEIR TOTAL LENGTHS 
(MM) AND LOCALITIES 


C. (S.) cuborientalis 2 6.8 Cuba 

C. (S.) curassavica & 7.0; 292 Curacao 

C. (S.) hummelincki 6 4.75.09 8:5 Aruba 

C. (S.) motasi 6 6:85 9100 Haiti 

C. (S.) orghidani 28:0 Cuba 

C. (S.) parapotamica ovis. 931692401 Jamaica 

C. (S.) salpiscinalis 6 5:6; Lares Haiti 

C. (S.) sbordonii 6, 2 90 Vera Cruz, Mexico 
C. (S.) specus 6) 18.0; 2 19:8 Cuba 

C. (S.) univam 2 10.0 Venezuela 


10 pectinate marginal spines; low triangular ridge present. Pleotelson evenly 
tapering to notched apex; angle of apex about 90°. 


RECORDS Oriente Province, Cuba, interstitial in river alluvia. 


Cyathura (Stygocyathura) curassavica Stork, 1940 
Figures 11E,F; 12 


DIAGNOSIS 6 7.0 mm, 2? 9.2 mm. Pereopod 1, propodal palm gently con- 
vex, bearing about 10 pectinate spines. Pleotelsonic margins in anterior two- 
thirds subparallel, tapering gently to finely notched apex; angle of apex less 
than 90°. 


RECORDS Curacao, from pits and wells. 


Cyathura (Stygocyathura) hummelincki Botosaneanu and Stock, 1982 
Figure 11G,H 


DIAGNOSIS ¢ 4.75 mm, ? 8.5 mm. Pereopod 1, propodal palm gently sinu- 
ate, bearing 11-23 marginal pectinate spines. Pleotelsonic margins faintly 
concave in midregion, posterior margin evenly convex, apex with slight 
notch. 


RECORDS Aruba, in pits, wells, and temporary water sources. 


36 ANTHURIDEA * ANTHURIDAE 


Figure 12. Cyathura 
(Stygocyathura) curassavica: 


pleon. 


Cyathura (Stygocyathura) motasi Botosaneanu and Stock, 1982 
Figure 111,J 


DIAGNOSIS <6 6.8mm, 2 10.0 mm. Pereopod 1, propodal palm gently sinu- 
ate, bearing 11—18 marginal pectinate setae. Pleotelsonic margins tapering 
evenly to slightly notched apex; angle of apex less than 90°. 


RECORDS Haiti, from wells. 


Cyathura (Stygocyathura) orghidani Negoescu Vladescu, 1983 
Figure 13A,B 


DIAGNOSIS ¢ 8.0 mm. Pereopod 1, propodal palm slightly convex, bearing 
about 11 marginal pectinate setae. Pleotelson, angle of apex obtuse, with 
small notch. 


RECORDS Pinar del Rio Province, Cuba, from freshwater lake in cave. 


Cyathura (Stygocyathura) parapotamica Botosaneanu and Stock, 1982 
Figure 13D,E 


DIAGNOSIS 2 4.1 mm (ovig. ? 3.6 mm). Pereopod 1, propodal palm sinu- 
ate, bearing eight marginal pectinate setae. Pleotelson gently tapering to 
notched apex; angle of apex about 90°. 


RECORDS Jamaica, from river alluvia. 


Figure 13. Cyathura (Stygocyathura) orghidani (from Negoescu, 1983): A, telson; B, 
pereopod 1, 2. Cyathura (Stygocyathura) sbordonii: C, pereopod 1, ° (from Argano, 
1971). Cyathura (Stygocyathura) parapotamica: D, telson; E, pereopod 1, ?. Cyathura 
(Stygocyathura) salpicinalis: F, pereopod 1, 2; G, telson. Cyathura (Stygocyathura) 
specus: H, 2; I, telson and uropods; J, pereopod 1, 2. Cyathura (Stygocyathura) 
univam: K, telson. 


38 ANTHURIDEA * ANTHURIDAE 


Cyathura (Stygocyathura) salpiscinalis Botosaneanu and Stock, 1982 
Figure 13F,G 


DIAGNOSIS 6 5.6 mm, 2 7.3 mm. Pereopod 1, propodal palm sinuate, 
bearing up to 15 marginal pectinate setae, and with distinct triangular ridge. 
Pleotelson gently tapering, with slight apical eminence. 


RECORDS Haiti, from alluvia of lake. 


Cyathura (Stygocyathura) sbordonii Argano, 1971 
Figure 13C 


DIAGNOSIS. 6 and 2 9.0 mm. Pereopod 1, propodal palm convex, bearing 
up to 16 marginal pectinate setae. Pleotelson with angle of apex obtuse. 


RECORDS Vera Cruz, Mexico, from freshwater in cave. 


Cyathura (Stygocyathura) specus Bowman, 1965 
Figure 13H-J 


DIAGNOSIS 6 18.0 mm, 2 19.8 mm. Pereopod 1, propodal palm almost 
straight, bearing up to 15 marginal pectinate setae. Pleotelson with angle of 
apex obtuse. 


RECORDS Las Villas Province, Cuba, from freshwater lake in cave. 


Cyathura (Stygocyathura) univam Botosaneanu, 1983 
Figure 13K 


DIAGNOSIS @ 10.0 mm. Pereopod 1, propodal palm gently convex, bearing 
22 marginal pectinate setae. Pleotelson posteriorly broadly rounded, apex 
emarginate. 


RECORDS Peninsula de Morocoy, Venezuela, from phreatic water in cave. 


Eisothistos Haswell, 1884 


DIAGNOSIS Mouthparts forwardly produced, tailfan spiny, indurate. Eyes 
present, larger and with more ommatodia in ¢ than in 9. Pereonites some- 
times elongate. Mandible with strong incisor, reduced lamina dentata, palp 
and molar lacking. Maxilliped lacking endite, palp slender, of three to five 


Eisothistos teri 39 


articles. Pereopods 1-3 not subchelate, propodi relatively elongate, mini- 
mally expanded. Pleopod 1 2, rami fused, together forming operculum; d 
rami separate. Pleonites free, short, longer in d than in 2. Telson lacking 
statocysts. 


REMARKS Wagele (1979) first recorded species of Evzsothistos preying on ser- 
pulid polychaete worms in their tubes. 

The genus contains about 12 species in the Pacific, Indian Ocean, Carib- 
bean, Antarctic, and Mediterranean. 


Key to species of Eisothistos 


1. @ telson with middorsal spines; d pereopod | propodal palm with 19 
Gi AURSOUES tee nie eile Suagece ane 2 ke ooo 1s eeu Gm ata aN teri 

2 telson lacking middorsal spines; d pereopod | propodal palm with 
MRS SIOIINE SR ls Chia os spoc sens Meat: «she ieccusen a, eee ie ras oo ale anegey ae petrensis 


Eisothistos petrensis Kensley, 1984 
Figure 14A—E 


DIAGNOSIS @: 4.0 mm. Antennular flagellum of six articles. Antennal 
flagellum of six articles. Telson posteriorly faintly bilobed, margin strongly 
serrate; faint anterior middorsal ridge; middorsally unarmed. Pereopod 1, 
propodal palm unarmed. Pleopod | rami fused for '/12 of length. d: 2.0 mm. 
Antennular flagellum of eight articles. Antennal flagellum of six articles. Per- 
eopod | propodal palm with 11 fringed spines. Telson narrower than in &, 
with middorsal ridge running almost entire length. 


RECORDS Carrie Bow Cay, Belize, 0.1—-36 m; Looe Key, Florida, 5—6 m; 
Turks and Caicos Islands, 1.0 m; St.Thomas, U.S. Virgin Islands, 7-10 m. 


Eisothistos teri Kensley and Snelgrove, 1987 
Figure 14F—H 


DIAGNOSIS @: 3.2 mm. Basal antennular peduncle article bearing triangu- 
lar apically rounded laminate process; flagellum of seven articles. Antennal 
flagellum of six articles. Mandible with biserrate lamina dentata. Pereopod 1 
propodal palm unarmed. Pleopod | rami fused for */+ of length. Uropodal 
exopod with one or two strong slightly recurved spines on dorsal surface. 


40 


ANTHURIDEA * ANTHURIDAE 


i 


: SYK Bee: 


Figure 14. Eisothistos petrensis: A, 2; B, uropodal sympod and endopod; C, 
uropodal exopod; D, pleopod 1; E, pereopod 1, ¢. Eisothistos teri: F, telson and 


uropod; G, pleopod 1; H, pereopod 1, d. 


Telson with eight or nine slightly recurved middorsal teeth becoming longer 
posteriorly. d: 2.0 mm. Antennular flagellum of eight articles. Antennal 


Haliophasma curri 4] 


flagellum of six articles. Pereopod 1 propodal palm with 19 or 20 fringed 
spines. Uropodal exopod lacking dorsal teeth. Telson lacking middorsal 
teeth; posterior margin incised into 12 acute or narrowly rounded teeth. 


RECORDS’ Barbados, in Madracis mirabilis coral, 9-15 m. 


Haliophasma Haswell, 1881 


DIAGNOsIS_ Eyes present. Integument often indurate, with scattered pitting. 
Antennular flagellum usually of two articles. Antennal flagellum of 4—7 arti- 
cles. Mandibular palp of three articles. Maxillipedal palp of two articles, 
article 2 smaller than article 1. Pereopod 1, propodus expanded. Pereopods 
4—7, carpi roughly rectangular. Pleopod | exopod operculiform. Pleonites 1— 
5 short, fused; pleonite 6 usually demarked from telson. Latter with two 
basal statocysts. d often with more elongate form than ?. Antennular 
flagellum multiarticulate. Eyes larger. 


Key to species of Haliophasma 


1. Telson posteriorly narrowly rounded; dactylus of pereopod | dentate 
sige MN NE AE ey Sees ence Dene Cara anes ey Par eee we pee valeriae 


Haliophasma curri Paul and Menzies, 1971 
Figure 15A—C 


DIAGNOSIS @ 7.0 mm. Antennular flagellum of three articles. Antennal 
flagellum of five articles. Mandibular palp article 3 with five spines. Max- 
illipedal palp, terminal article small, set obliquely at distolateral angle of 
article 1. Pereopod 1, carpus triangular, posterodistal margin crenulate, dis- 
tally rounded; propodal palm crenulate, with low rounded proximal lobe. 
Pleonite 6 dorsally demarked, narrow, with middorsal point in posterior 
margin. Uropodal exopod elongate, outer margin sinuate, dentate; endopod 
ovate, distally narrowed, length twice greatest width, outer margin dentate. 
Telson parallel sided, posterior margin broadly rounded. 


RECORDS Off Venezuela, 95 m; Culebra Island, Bay of Panama, intertidal. 


42 ANTHURIDEA ¢ ANTHURIDAE 


Figure 15. Haliophasma curri: A, 2; B, pereopod 1; C, uropod. Haliophasma valeriae: 
D, 2; E, pereopod 1; F, uropod. 


Haliophasma valeriae Paul and Menzies, 1971 
Figure 15D-F 


DIAGNOSIS 2 6.5 mm. Body long and slender. Antennular flagellum of 


Malacanthura 43 


three articles. Antennal flagellum of six articles. Mandibular palp article 3 
with four spines. Pereopod 1, carpus triangular, tipped with acute tooth; 
propodus elongate, palm with about seven teeth, five fringed spines on mesial 
surface; unguis of dactylus strongly flexed; margin of dactylus with three 
strong triangular teeth. Uropodal exopod elongate, apically acute, margins 
serrate; endopod length little more than twice greatest width, outer margin 
serrate, apex acute. Telson elongate-elliptical, apically narrowly rounded; 
strong middorsal longitudinal rounded ridge running almost entire length. 


RECORDS Off Venezuela, 95 m. 


REMARKS Wagele (1981) made this species the type of his new genus Ne- 
manthura, based primarily on the elongate form of the body and appendages. 
Haliophasma irmae Paul and Menzies, 1971, from the same locality as the 
above species, is probably the same species. 


Licranthura Kensley and Schotte, 1987 


DIAGNOSIS Serrate process on antennal peduncle article 3. Mandibular 
palp of three articles; molar lacking. Maxillipedal endite short. Pereopod 1 
larger than pereopods 2 and 3. Pereopods 4—7, carpi triangular. Pleonites 
short, free. Pleopod 1, both rami forming operculum. Pleotelson lacking 
statocysts. 


Licranthura amyle Kensley and Schotte, 1987 
Figure 16 


DIAGNOSIS @ 3.8 mm. Eyes small, pigmented. Antennular flagellum of 
three articles. Antennal peduncle article 3 with lamellar expanded process, 
serrate on mesial margin; flagellum of six articles. Maxillipedal palp of five 
articles; very short endite. Pereopod 1, propodal palm unarmed. Uropodal 
and telsonic margins serrate. 


RECORDS Carrie Bow Cay, Belize, 0—25 m, in coral rubble. 


Malacanthura Barnard, 1925 


DIAGNOSIS. Eyes present. Mandibular palp of three articles. Maxillipedal 
palp of three articles, terminal article usually broadly ovate. Pereopod 1, 
propodus expanded. Pereopods 4—7 with carpi roughly rectangular. Pleopod 


44 ANTHURIDEA * ANTHURIDAE 


Figure 16. Licranthura amyle: A, 2; B, antennule and antenna; C, telson and 
uropod. 


1, exopod operculiform. Pleonites 1—5 short, fused; pleonite 6 dorsally de- 
marked. Pleotelson with two basal statocysts. 


Mesanthura 45 


Malacanthura caribbica Paul and Menzies, 1971 
Figure 17 


DIAGNOSIS @ 27.1 mm. Integument moderately indurate. Antennular 
flagellum of seven articles. Antennal flagellum of four articles. Mandibular 
palp, article 3 with comb of 11 spines. Maxillipedal palp, terminal article 
broadly ovate, penultimate article with row of seven spines on mesial margin. 
Pereopod 1 propodus expanded, palm straight, with few spines on mesial 
margin. Uropodal exopod barely reaching base of endopod, narrow, apically 
acute, outer margin sinuate, serrate; endopod set obliquely on sympod, mar- 
gin serrate, apically acute. Telson lanceolate, apically narrowly rounded, 
with strong longitudinal middorsal carina. 


RECORDS Off Venezuela, 95 m; off Colombia, 42—44 m. 


REMARKS Malacanthura cumanensis Paul and Menzies, 1971, described from 
the same locality as M. caribbica, was shown to be the latter species (Kensley, 
1980). 


Mesanthura Barnard, 1914 


DIAGNOSIS @: Dorsal integument with (usually) species-specific pigment 
pattern; pigment persistent in alcohol. Mandibular palp of three articles, 
terminal article with row of spines, number of which specific for species. 
Maxilliped with endite either very reduced or absent; palp of three articles, 
with terminal article usually about half length of penultimate article, suture 
transverse. Pereopod 1, propodus expanded, palm often with step. Pereopods 
2 and 3, propodi not expanded. Pereopods 4—7, carpi roughly triangular, 
with anterior margin shorter than posterior margin. Pleonites 1—5 fused, 
pleonite 6 dorsally demarked. Pleopod 1, exopod operculiform. Telson with 
two basal statocysts. d: Eyes larger than in 2. Antennular flagellum of sel- 
dom more than 10 articles bearing numerous aesthetascs. Mouthparts, espe- 
cially body of mandible, reduced. Pereopod 1, propodus bearing dense band 
of spines on mesial surface near palm. Pigment pattern more diffuse than in 
2, extending onto ventral surface. 


REMARKS Méesanthura is a relatively large genus of about 30 species, re- 
corded from most tropical and temperate seas, in shallow habitats. The 
males of few species have been recorded; by themselves, males are difficult to 
identify as the dorsal pigment pattern characteristic of the female breaks 
down and spreads onto the ventrum. 


Figure 17. Malacanthura caribbica: A, 2; B, °, lateral view; C, mandible; D, 
maxilliped; £, uropod; F, pleopod 1. 


Mesanthura bivittata 47 


Key to species of Mesanthura (° only) 


1. Pigment in tiny evenly scattered chromatophores over body; cephalon 
with solid patch of pigment; mandibular palp article 3 with seven 
SOUT COMI. co UMMeR a ceo His Hees og eee tn cst ot PRE cS punctillata 

Pismentinot evenly scattered over body = ..q.cn2... 40 See oes See 2s Z 


2. Pereonites 4 and 5 with patch of pigment, sometimes with unpigmented 
areammommiccdle Of pateliterc.. 0 <=/. sgt ee etce «shape apie s 2 sm ose Skt 3 
Pereonites 4 and 5 lacking fairly solid patch of pigment ............ 6 


3. Pereonites 4 and 5 lacking unpigmented area in pigment patch; five 
transverse lines on pigment on pleon; mandibular palp article 3 with 


STAG SPLINES gee sehoee ae ete, Ee. o's =, ae does Wats Sak tome eer Se paucidens 
Pereonites 4 and 5 with unpigmented area in pigment patch; pleon 
lackingjtransverse pigmentdines ...)9. «sens. os Species went hea 4 


4. Pigment in obvious double longitudinal bands on pereon and pleon; 
mandibular palp article 3 with nine spines ................ bivittata 
Picmentnotin obvious double bands. . 2 njcui-l2 soc nste ees Bee ey oo D 


5. Unpigmented area in middle of pigment patch of pereonites 1—3; 


mandibular palp article 3 with 10 spmes: 4......... 55. sa... .- pulchra 
No unpigmented area in middle of pigment of pereonites 1—3; 
mandibular palp article S"with eight spines”... 2.22242...» ©- looensis 


6. Pigment of pereon in fine reticulate lines; mandibular palp article 3 
WAGE SIXUS[ONMIES®  </o).<cpate NMC a 10 ve =.+ « ahtnee name» 6 = ees Aeon Sete reticulata 
Pigment of pereon not in fine reticulate lines; mandibular palp article 3 
SAL PlaN TOU SUING Ses Se cotedar hates = <> + het nears) cis, MaaPe ae arae oye 95, wv eg @ 7 


7. Pigment in more or less complete rings on pereonites 1-6 ..... hopkinsi 
Pigment in strong transverse posterior bars on pereonites 4—7 . . fasciata 


Mesanthura bivittata Kensley, 1987a 
Figures 18A, 20A—D 


DIAGNOSIS. 6 5.2 mm, ovigerous 2? 7.8 mm. Pigment in obvious double 
longitudinal bands on pereon and pleon. Mandibular palp article 3 with nine 
spines. Maxilliped lacking endite. Pereopod 1, propodal palm with rounded 
lobe. 


RECORDS ‘Twin Cays, Belize, under red mangroves, 1—2 m. 


Mesanthura hopkinsi; D, 


BI 


Mesanthura bivittata; B, Mesanthura fasciata; C 


3 


Figure 18. A 


Mesanthura looensis. 


Mesanthura fasciata 49 


e 
wi) 
qo 
rd 


ed 
ene 


Se 
wes 
vee 

: 


Figure 19. A, Mesanthura paucidens; B, Mesanthura pulchra; C, Mesanthura punctillata; 
D, Mesanthura reticulata. 


Mesanthura fasciata Kensley, 1982 
Figures 18B, 20E—H 


DIAGNOSIS @ 4.5 mm. Pigment in triangular patch on head, open irregular 
rings on pereonites 1—3; pereonites 4—7 with strong transverse posterior bar; 


Figure 20. Mesanthura bivittata: A, mandible; B, maxilliped; C, pereopod 1; D, 
uropodal exopod. Mesanthura fasciata: E, mandible; F, maxilliped; G, pereopod 1; 
H, uropodal exopod. Mesanthura hopkinsi: I, mandible; J, maxilliped; K, pereopod 1; 
L, uropodal exopod. 


Mesanthura paucidens 51 


five bars on pleon. Mandibular palp article 3 with four spines. Maxilliped 
with very reduced endite. Pereopod 1, propodal palm with step. 


RECORDS Looe Key, Florida, 5—6 m; Cozumel, Mexico; Jamaica; Carrie 
Bow Cay, Belize, 0.2—6 m. 


Mesanthura hopkinsi Hooker, 1985 
Figures 18C, 20I-L 


DIAGNOSIS ¢ 2.4 mm. Pigment in triangle on cephalon; in irregular rings 
on pereonites 1—6; four transverse bars on pleon. Mandibular palp article 3 
with four spines. Maxilliped lacking endite. Pereopod 1, palm of propodus 
lacking step. 


RECORDS Looe Key, Florida, 0.5 m; Florida Middlegrounds, Gulf of Mex- 
Ico, 59°m: 


Mesanthura looensis Kensley and Schotte, 1987 
Figures 18D, 21A—D 


DIAGNOSIS 10.0 mm. Pigment in solid patches on cephalon, pereonites, 
pleon, uropods and telson; pereonites 4—6 with open central area in patch. 
Mandibular palp article 3 with eight spines. Maxilliped lacking endite. Per- 
eopod 1, propodal palm with step. 


RECORDS Looe Key, Florida, | m. 


Mesanthura paucidens Menzies and Glynn, 1968 
Figures 19A, 21E-I 


DIAGNOSIS @:6.6 mm. Pigment in roughly rectangular to ovate patches on 
cephalon and pereonites; in five transverse lines connected laterally on pleon. 
Mandibular palp article 3 with six spines. Maxilliped with short narrow en- 
dite. Pereopod | propodal palm with step. d: 6.4 mm. Antennular flagellum 
of seven articles. Pigment more diffuse than in 2 but retaining five pleonal 
bars. 


RECORDS Looe Key, Florida, 5—6 m; Carrie Bow Cay, Belize, 15.2 m; Pu- 
erto Rico, intertidal; Jamaica. 


52 ANTHURIDEA * ANTHURIDAE 


o 


Figure 21. Mesanthura looensis: A, mandible; B, maxilliped; C, pereopod 1; D, 
uropodal exopod. Mesanthura paucidens: E, mandible; F, maxilliped; G, pereopod 1, 
2; H, pereopod 1, 6; J, uropodal exopod. Mesanthura pulchra: J, mandible; K, 
maxilliped; L, pereopod 1, d; M, pereopod 1, 2; N, uropodal exopod. 


Mesanthura pulchra Barnard, 1925 
Figures 19B, 21J—N 


DIAGNOSIS 2: 9.3 mm. Pigment pattern in roughly rectangular patches on 
cephalon and pereonites 1—6, with open oval middorsal area in patch. Man- 
dibular palp article 3 with 10 spines. Maxilliped lacking endite. Pereopod 1, 
propodal palm with step. d: 5.4 mm. Antennular flagellum of eight articles. 


Minyanthura corallicola 53 


RECORDS Egmont Key, Florida, 18.3—36.6 m; Looe Key, Florida, 0.5—12 
m; Dry Tortugas; Turks and Caicos Islands, | m; Puerto Rico, intertidal to 
1.5 m; Carrie Bow Cay, Belize, intertidal to 1.5 m; St. Thomas and St.John’s, 
U.S. Virgin Islands; Cozumel, Mexico. 


REMARKS Menzies and Glynn (1968) recorded this species as M. decorata 
from Puerto Rico, while Menzies and Kruczynski (1983) recorded it as M. 
floridensis. 


Mesanthura punctillata Kensley, 1982 
Figures 19C, 22A—F 


DIAGNOSIS 2: 6.4 mm. Pigment in solid patch between eyes, rest of pig- 
ment on pereon and pleon with chromatophores scattered, diffuse, not in any 
regular pattern. Mandibular palp article 3 with seven spines. Maxilliped 
lacking endite; terminal palp article semicircular. Pereopod | propodal palm 
with step. d: 4.5 mm. Antennular flagellum of 10 articles. 


RECORDS ‘Turks and Caicos Islands, 1 m; Carrie Bow Cay, 0.2—20 m. 


Mesanthura reticulata Kensley, 1982 
Figures 19D, 22G—J 


DIAGNOSIS { 6.1 mm. Dorsal pigment pattern a network of 
chromatophores arranged in fine lines. Mandibular palp article 3 with six 
spines. Maxilliped lacking endite. Pereopod 1 propodal palm with step. 


RECORDS Carrie Bow Cay, 10—24 m. 


Minyanthura Kensley, 1982 


DIAGNOSIS Mandible lacking molar and palp. Maxillipedal palp of five ar- 
ticles; large apically acute endite present. Pereopods 4—7, carpi rectangular. 
Pleopod 1, both rami forming operculum. Pleonites 1—5 fused; pleonite 6 
fused with telson, not dorsally demarked. Telson with two basal statocysts. 


Minyanthura corallicola Kensley, 1982 
Figure 23 


DIAGNOSIS @: 1.7 mm. Antennular flagellum of one article. Antennal 


Figure 22. Mesanthura punctillata: A, mandible, 2; B, mandible 6; C, maxilliped; D, 
pereopod 1, 2; E, pereopod 1, 6; F, uropodal exopod. Mesanthura reticulata: G, 
mandible; H, maxilliped; J, pereopod 1; J, uropodal exopod. 


Minyanthura corallicola 55 


Figure 23. Minyanthura corallicola: A, 2; B, telson and uropod; C, pleopod 1; D, 
mandible; E, maxilliped; F, pereopod 1. 


flagellum of four articles. Pereopod 1, propodus somewhat inflated. Uropo- 
dal rami and posterior telsonic margin serrate. d: 1.3 mm. Eyes larger than 


in 2. Antennular flagellum of three articles. Antennal flagellum of two 
articles. 


RECORDS Carrie Bow Cay, Belize, 6-24 m; Barbados, 9-15 m; Jamaica. 


56 ANTHURIDEA * ANTHURIDAE 


Pendanthura Menzies and Glynn, 1968 


DIAGNOSIS 2: Integument with some red-brown dorsal pigmentation. Eyes 
small, pigmented. Antennular flagellum of two articles. Antennal flagellum 
of one article. Mandibular palp of single reduced article; incisor, molar, and 
lamina dentata present. Maxillipedal palp of single broad article; small tri- 
angular endite present. Pereopod 1, propodus expanded. Pereopods 4-7, 
carpi short, triangular, lacking free anterior margin. Pleonites 1—5 very 
short, fused, 6 fused with telson. Pleopod 1, exopod operculiform. Telson 
basally broad, with two statocysts at about midlength. d: Antennular ped- 
uncle of four articles. Antennal flagellum of one article. Pereopod 1, pro- 
podus with dense clump of spines on mesial surface. Eyes only slightly larger 
than in @. 


REMARKS The genus comprises three species, two from the Caribbean and 
one from the Pacific, all of which have been taken from shallow coral reefs. 


Key to species of Pendanthura 


1. Dorsal pigmentation over entire body; pereopod 1, propodal palm with 

rounded bey 2 nak seeps Sees mot ee eee tanaiformis 
Dorsal pigmentation on cephalon, pereonite 2, and pleon; pereopod 1, 

propedal palmulacking rounded lobe fonts = 2 se hendleri 


Pendanthura hendleri Kensley, 1984 
Figure 24A—E 


DIAGNOSIS 2: 3.3 mm. Dorsal pigmentation limited to cephalon, pereonite 
2, and very short pleon. Reduced mandibular palp with two setae. Pereopod 
1, propodus expanded, palm gently convex, lacking rounded lobe, with four 
spines on mesial surface near palmar margin. d: 2.8 mm. Pigmentation as in 
?. Pereopod 1, propodal palm gently convex, lacking rounded lobe. 


RECORDS Carrie Bow Cay, Belize, 9-23 m; Twin Cays, Belize, 0Q—2 m; Pan- 
ama, 30 m. 


Pendanthura tanaiformis Menzies and Glynn, 1968 
Figure 24F-H 


DIAGNOSIS Ovigerous 2: 2.9 mm. Dorsal pigmentation a dense red-brown 


Figure 24. Pendanthura hendleri: A, 2; B, mandible; C, maxilliped; D, pereopod 1, 
3; E, pereopod 1, 2. Pendanthura tanaiformis: F, pereopod 1, 2; G, 2; H, mandible. 


58 ANTHURIDEA * HYSSURIDAE 


reticulation over entire dorsum. Mandibular palp with one seta. Pereopod 1, 
propodus expanded, palm with rounded lobe, mesial surface with six spines 
near palmar margin. d: 2.8 mm. Pigmentation as in 2. Propodal palm with 
rounded lobe. 


RECORDS’ Bermuda; Carrie Bow Cay, Belize, intertidal to 1 m; Puerto Rico, 
intertidal; Cozumel, Mexico. 


REMARKS Kensley (1984) characterized this common species of the interti- 
dal of the reef crest as stress-tolerant, breeding throughout the year. 


Skuphonura Barnard, 1925 


DIAGNOSIS Antennal flagellum of single article. Cephalon with midventral 
toothlike process at base of mouthparts. Mandibular palp of three articles. 
Maxillipedal palp of three articles; endite lacking. Pereopod 1, propodus ex- 
panded. Pereopods 4—7, carpi triangular. Pleopod 1, exopod operculiform. 
Pleonites 1—5 fused; pleonite 6 dorsally demarked. Pleotelson with two basal 
statocysts. 


Skuphonura laticeps Barnard, 1925 
Figure 25 


DIAGNOSIS 6 6.0 mm. Cephalon wider anteriorly than posteriorly, with 
anterolateral lobes extending well beyond rostrum. Antennular flagellum of 
two articles. Pereonite 1 with strong midventral forwardly directed tooth. 
Pereopod 1, carpus with posterodistal angle produced into triangular lobe; 
propodus expanded, palmar margin proximally convex, numerous setae on 
mesial surface. Pleopod 2, copulatory stylet of endopod reaching beyond 
rami. Pleonite 6 dorsally demarked, posterior margin middorsally incised. 
Uropodal exopod ovate, with distal notch; endopod length slightly more than 
basal width. Pleotelson widest at midlength, posteriorly narrowly rounded, 
with broadly rounded longitudinal raised area. 


RECORDS St. Thomas, U.S. Virgin Islands, 8—40 m. 


Family Hyssuridae Wagele, 1981 


DIAGNOSIS. Maxillipedal palp usually of five articles; endite present. Per- 
eopods 1—3 subsimilar, often all three pairs subchelate. Pleopod | similar to 


Figure 25. Skuphonura laticeps: A, 3; B, cephalon and pereonite 1, lateral view; C, 
antennule, d; D, pereopod 1, d; £, antenna; F, maxilliped. 


60 ANTHURIDEA * HYSSURIDAE 


following pleopods, not operculiform. Pleonites 1—5 freely articulating, rela- 
tively elongate. Pleotelson lacking statocysts. 


Key to genera of Hyssuridae 


1. Pleotelson not covered by uropodal exopods; maxillipedal rami basally 
EEE) a. Son SE sists aS he ed ES ey, Send ol SPs Ae Sree ae Kupellonura 

Pleotelson covered completely by uropodal exopods; maxillipedal rami 
basally fused) ic) c-f5.1.) errs cose oar te ea ae cess ea tor earns Xenanthura 


Kupellonura Barnard, 1925 


DIAGNOSIS Mandibular palp of three articles. Maxillipedal palp of five arti- 
cles; large endite present. Pereopods 1—3 similar, propodi somewhat ex- 
panded. Pereopods 4—7, carpi triangular. Pleonites 1—5 elongate, free, sube- 
qual. Pleopods 1-5 similar. 


Kupellonura imswe (Kensley, 1982) 
Figures 26, 27 


DIAGNOSIS @: 3.4 mm. Eyes present. Antennular flagellum of four articles. 
Antennal flagellum of seven articles. Maxillipedal endite reaching to base of 
article 3. Pereopods 1-3 similar; pereopod 1, carpus posterodistally acute; 
propodus expanded, palm straight, unarmed; carpus triangular, free anterior 
margin shorter than posterior. Pleonite 6 with middorsal incision in posterior 
margin. Uropodal exopod, outer margin serrate, distally narrowly rounded; 
endopod length twice greatest width. Telson widest at midlength, 
posterolateral margin serrate, apically broadly rounded. d: 2.7 mm. Eyes 
larger than in 2. Mouthparts reduced. Pereopod 1, propodal palm armed 
with row of eight fringed spines. 


RECORDS Carrie Bow Cay, Twin Cays, Glover’s Reef, Belize, 0—6.0 m; 
Montego Bay, Jamaica, | m. 


Xenanthura Barnard, 1925 


DIAGNOSIS Mandibular palp of single article. Maxillipedal rami fused; ar- 
ticulations of palp articles obscure. Pereopods 1—3 similar. Pereopods 4—7, 
carpi triangular. 


Figure 26. Kupellonura imswe: A, 2°; B, cephalon and pereonite 1, 3; C, maxilliped; 


D, pereopod 1, ; E, pereopod 1, 3. 


62 ANTHURIDEA * HYSSURIDAE 


Figure 27. Kupellonura imswe: A, 2 cephalon, dorsal view; B, posterior pleon, dorsal 
view; C, pleon, ventral view; D, cephalon and pereonites | and 2, lateral view. 


Xenanthura brevitelson Barnard, 1925 
Figure 28 


DIAGNOSIS 2: 4.0 mm. Ommatidia of eyes in longitudinal row of three or 
four groups. Antennular flagellum of three articles. Antennal flagellum of 
three articles. Mandibular palp of single short article bearing single seta. 
Maxillipedal rami basally fused, palp of three or four obscurely separated 
articles. Pereopods 1—3 similar, carpus of pereopod 2 triangular, with strong 
triangular projecting lobe posterodistally; propodi expanded, palm incised 
into several rounded lobes. Uropodal exopods circular, overlapping dorsally 
and covering telson; endopod projecting beyond exopods ventrally, mesial 
margin with step, distally rounded. Telson shorter than uropodal exopod, 
posterior margin truncate to faintly concave. 6: 3.5 mm. Antennular 
flagellum of seven articles. 


RECORDS Off Georgia, 20-145 m; off Florida, 8-10 m; Turks and Caicos 
Islands, 1 m; St.Thomas, U.S. Virgin Islands, 50-60 m; Gulf of Mexico. 


Figure 28. Xenanthura brevitelson: A, @; B, pereopod 1; C, uropodal endopod; D, 


telson; E, uropodal exopod. 


64 ANTHURIDEA * PARANTHURIDAE 


Family Paranthuridae Menzies and Glynn, 1968 


DIAGNOSIS Mouthparts together forming somewhat elongate cone adapted 
for piercing and sucking. Mandible with styliform incisor, lacking lamina 
dentata and molar. Maxilla slender, styliform, distally serrate. Maxilliped 
elongate; number of palp articles usually reduced. Pereopod 1, or pereopods 
1-3 subchelate. Pleopod 1 exopod operculiform. Pleonites short, free or 
fused. Pleotelson with single basal statocyst, or lacking statocyst. 


Key to genera of Paranthuridae 


1. Pereopod»7 lacking in‘adult. s/o. 92) sashes sao dee-.> so ee eee 2 
Pereopod. 7 present inadult: 2. a. %ielas om gees ole sd tok cates oe eee 3 
2: EVES (PLESEME (6.05. jn Sees. Sao, seeks cobs tonlere sale Sie eins etd Ss < fete Colanthura 
Byes absent. is... ci eee Seem mL ans oo Ee Curassanthura 


3. Antennular and antennal flagellum of more than 10 
anticles: 37.5 ae: dae ee eae os Shee Meee Misia eee eee Accalathura 


4. Antennal flagellum a single (rarely two or 3) flattened article; 


maxilhpedal palp of onevor twetartcles’ 219. ..210025 os Paranthura 
Antennal flagellum of seven articles; maxillipedal palp of three articles 
eR Te ere ate Lerner 2. Sa GOs RRS OS cee ks Virganthura 


Accalathura Barnard, 1925 


DIAGNOSIS Eyes present. Antennular and antennal flagella multiarticulate, 
each of more than 10 articles. Mandibular palp of three articles. Maxillipedal 


Key to species of Accalathura 


1. Uropodal exopod elongate-narrow; endopod length twice basal width 
crenulata 


Uropodal exopod ovate, apically subacute; endopod length 1.5 times 
basal widthy (iho este fac siecnedenn eee eh Nera Chane ene tate ketene setosa 


Colanthura tenuis 65 


palp of two articles, endite almost reaching end of palp. Pereopod | subche- 
late, propodus inflated, larger than pereopods 2 and 3. Pereopods 4—7, carpi 
with anterior and posterior margin subequal. Pleonites free, short. Pleopod 1, 
exopod operculiform. Telson with single statocyst. 


Accalathura crenulata (Richardson, 1901) 
Figure 29A—D 


DIAGNOSIS 2: 16.0 mm. Antennular flagellum of about 26 articles. Anten- 
nal flagellum of about 18 articles. Uropodal exopod narrow, parallel sided. 
Telson apically subacute. d: 15.0 mm. Pleopod 2, copulatory stylet of endo- 
pod apically acute, with subapical “heel.” 


RECORDS Off North Carolina, 30 m; off Georgia, 20 m; Cuba; Puerto Rico, 
intertidal; Cozumel, Mexico; Carrie Bow Cay, Twin Cays, Belize, intertidal 
to 6 m; west coast of Florida, Gulf of Mexico, 55 m. 


Accalathura setosa Kensley, 1984 
Figure 29E—H 


DIAGNOSIS 2: 8.5 mm. Antennular flagellum of 11 articles. Antennal 
flagellum of 13 articles. Uropodal rami, margins closely setose; exopod ovate, 
outer margin sinuate, apex subacute; endopod ovate, length 1.25 times great- 
est width. Telson apically rounded. d: 7.0 mm. Pleopod 2, copulatory stylet 
of endopod apically strongly bifid. 


RECORDS Carrie Bow Cay, Belize, intertidal to 0.5 m. 


Colanthura Richardson, 1902 


DIAGNOSIS Integument with minute squamae. Mandible lacking palp. 
Maxillipedal palp articles fused except for minute terminal article. Pereopod 
1 subchelate, propodus expanded. Pereopods 2 and 3 subchelate but smaller 
than pereopod 1. Pereopods 4—6, carpi rectangular. Pereonite 7 very short, 
pereopod 7 lacking. Pleotelson lacking statocyst. 


Colanthura tenuis Richardson, 1902 
Figure 30A—C 


DIAGNOSIS : 3.5 mm. Eyes present. Integument diffusely brown in color. 
Antennular flagellum of four articles. Antennal flagellum of single article. 


Figure 29. Accalathura crenulata: A, 2; B, uropodal sympod and endopod; C, 
uropodal exopod; D, telson. Accalathura setosa: E, 2; F, uropodal sympod and 
endopod; G, uropodal exopod; H, telson. 


Curassanthura bermudensis 67 


Pereopod 1, propodus with mesial surface bearing proximal row of six spines. 
Pleonites 1—5 short, fused, boundaries marked dorsally by folds. Telson pos- 
teriorly broadly rounded. ¢: 3.5 mm. Antennular flagellum of five articles. 


RECORDS Bermuda, intertidal to 0.5 m. 


Curassanthura Kensley, 1981 


DIAGNOSIS. Eyes lacking. Mandibular palp of three articles. Maxillipedal 
palp of five articles; short endite present. Pereopod | subchelate. Pereopods 
2-6 similar, carpi rectangular. Pereopod 7 lacking. Pleonites 1—5 free; 
pleonite 6 dorsally demarked. Pleopod 1, exopod operculiform. Pleotelson 
with single statocyst. Interstitial littoral forms. 


REMARKS Three species of this interstitial genus are known, two from the 
Caribbean, and one from the upper sublittoral gravels of a lava tunnel on 
Lanzarote, Canary Islands. 


Key to species of Curassanthura 


1. Telson with posterior fourth abruptly narrowed; uropodal exopod 
length) 49 umes ereatest width. 25). of ac2e scr o3 gas bermudensis 
Telson tapering, but posterior fourth not abruptly narrowed; uropodal 
exopod! lengih 2-amimies ereatest Widths. ~.. <--ee - +25. --4-- halma 


Curassanthura bermudensis Wagele, 1985 
Figure 30G,H 


DIAGNOSIS @ 3.0 mm. Pereopod | propodal length 2.5 times proximal 
width, palm with proximal strongly recurved hooklike tooth. Uropodal ex- 
opod slender, parallel sided, 4.5 times longer than greatest width. Telson 
constricted in posterior fourth. 


RECORDS Church Cave, Bermuda, in shore sediments. 


68 ANTHURIDEA * PARANTHURIDAE 


Figure 30. Colanthura tenuis: A, 2; B, uropod; C, pereopod 1. Curassanthura halma: 
D, @; E, telson; F, uropodal exopod. Curassanthura bermudensis: G, telson; H, 
uropodal exopod (from Wagele, 1985). 


Curassanthura halma Kensley, 1981 
Figure 30D-F 


DIAGNOSIS @ 2.3 mm. Pereopod 1, propodal length about 1.7 times proxi- 
mal width, palm with eight fringed spines and basal tridentate lobe. Uropo- 


Paranthura antillensis 69 


dal exopod triangular, length 2.5 times greatest width, shorter than sympod. 
Telson tapering but not abruptly narrowed in posterior fourth. 


RECORDS Curagao, in shore sediments 1.5 m above tide line; Bonaire, in 
shore sediments above tide line. 


Paranthura Bate and Westwood, 1868 


DIAGNOSIS. Eyes present. Antennular flagellum shorter than peduncle. An- 
tennal flagellum usually of single flattened setose article. Mandibular palp of 
three articles, article 3 with comb of spines. Maxillipedal palp of one or two 
articles; endite small to absent. Pereopod 1, propodus inflated, larger than 
that of pereopods 2 and 3. Pereopods 4—7, carpi rectangular. Pleonites short, 
more or less distinct. Pleopod 1, exopod operculiform. Telson lacking 
statocyst. 


REMARKS This is the largest of the paranthurid genera, with over 50 names 
in the literature. Many of these are poorly described. Species of Paranthura are 
common in the shallow waters of the temperate and tropical seas. 


Key to species of Paranthura 


1. Telson posteriorly truncate; uropodal exopod rectangular, margins 
SeMEALe ) Pee Se OMe ae ER on Sh oe Re Side ice infundibulata 
Telson posteriorly rounded; uropodal exopox ovate, margins entire .. 2 


Za wropodaltendoped longer than wide ...5.¢.,. 445-00: 6 s-e206-% +s fos 3 
Wropodal-endopad as lone as wide... ..4. digs. s wscace cise antillensis 


3. Uropodal exopod elongate-elliptical; telson parallel sided for half length 


Uropodal exopod ovate, outer margin sinuate; telson evenly elliptical 
SRS rite il eres Sete akon ek eS ahh Op eae bidnt ga rece A a barnardi 


Paranthura antillensis Barnard, 1925 
Figure 31A—F 


DIAGNOSIS 2 5.1 mm. Antennular flagellum of four articles. Mandibular 
palp, article 3 with five spines. Maxillipedal palp of single article three times 
longer than basal width; short endite present. Pereopod 1, propodus ex- 


Figure 31. Paranthura antillensis: A, 2; B, telson; C, uropodal exopod; D, uropodal 
sympod and endopod; E, pereopod 1, 2; F, pereopod 2, 2. Paranthura barnardi: G, 
2; H, telson; J, pereopod 1, 2; J, uropodal exopod; K, uropodal sympod and 
endopod. 


Paanthura infundibulata 71 


panded, palm bearing row of setae, mesial surface near palmar margin with 
convex flange and row of about 10 spines. Pereopod 2, propodal palm with 
five stout sensory setae. Pleonite 6 free, posterior margin bilobed. Uropodal 
exopod ovate, outer margin sinuous; endopod almost circular, as wide as 
long. Telson posteriorly rounded. 


RECORDS St. Johns, St. James, U.S. Virgin Islands, 32 m; Carrie Bow Cay, 
Belize, intertidal to 1.5 m. 


Paranthura barnardi Paul and Menzies, 1971 
Figure 31G—K 


DIAGNOSIS @ 6.0 mm. Mandibular palp article 3 with eight spines. Max- 
illipedal palp of single article, length four times basal width. Pereopod 1, 
propodal palm concave, with convex flange and row of about 17 spines on 
mesial surface. Uropodal exopod broadly ovate, apically subacute; endopod 
ovate, length about 1.5 times basal width. Telson evenly elliptical, apex 
evenly rounded. 


RECORDS Off Venezuela, 95 m. 


Paranthura floridensis Menzies and Kruczynski, 1983 
Figure 32A—E 


DIAGNOSIS @ 6.3 mm. Integument with sparse irregular pigmentation. 
Mandibular palp article 3 with eight spines. Maxillipedal palp of single arti- 
cle, length 3.5 times basal width. Pereopod 1, propodal palm with trans- 
parent flange and row of 10 setae on mesial surface. Uropodal exopod 
elongate-elliptical; endopod ovate, length 1.5 times greatest width. Telson 
posteriorly broadly rounded, parallel sided for about half its length. 


RECORDS Off Sanibel Island, Florida, Gulf of Mexico, 73 m. 


Paranthura infundibulata Richardson, 1902 
Figure 32F-J 


DIAGNOSIS @ 8.2 mm, 6 6.0 mm. Integument with red-brown pigmenta- 
tion; broad irregular patch between eyes running onto bases of antennules; 
pereonites | and 2 with anterior patches, remainder of pereonites with pos- 
terior patches; strong patch on uropodal exopod, endopod, and telson. Man- 
dibular palp article 3 with 11 or 12 spines. Maxillipedal palp of single article, 


Figure 32. Paranthura floridensis: A, 2; B, pereopod 1, 2; C, uropodal sympod and 
endopod; D, uropodal exopod; E, telson. Paranthura infundibulata: F, 2; G, telson; 
H, pereopod 1, 2; J, uropodal exopod; J, uropodal sympod and endopod. 


ASELLOTA ns 


length 2.5 times basal width. Pereopod 1, propodus with convex flange and 
row of more than 20 setae. Uropodal exopod elongate-rectangular, mesial 
and distal margins serrate; endopod roughly square, margins serrate. Telson 
parallel sided, posterior margin truncate. 


RECORDS Bermuda, 11—12 m; Carrie Bow Cay, Belize, intertidal to 1 m; 
Cozumel, Mexico; Venezuela. 


Virganthura Kensley, 1987b 


DIAGNOSIS’ Eyes present. Mandibular palp of three articles. Maxillipedal 
palp of three articles; endite present. Pereopods 1—3 subchelate, pereopod | 
larger than pereopods 2 and 3; pereopods 4—7, carpi rectangular. Pleopod 1, 
exopod operculiform. Pleonites 1—5 short, distinct; pleonite 6 dorsally de- 
marked. Telson with single statocyst. 


Virganthura crassa (Barnard, 1925) 
Figure 33 


DIAGNOSIS ? 6.8 mm. Antennular flagellum of three articles. Antennal 
flagellum of seven articles. Maxillipedal endite reaching distal margin of first 
palp article. Pereopod 1, propodal palm slightly concave, bearing seven 
spines. Uropodal exopod ovate, outer margin sinuate; endopod distally 
rounded, articulating obliquely on sympod. 


RECORDS USS. Virgin Islands, 30 m. 


Suborder Asellota Latreille, 1803 


DIAGNOSIS Antennules uniramous. Antennae uniramous, with scale in 
some families. Mandible usually with palp, but palp lacking in some groups. 
Pereopod 1 usually subchelate, sexually dimorphic in some groups. Coxae 
small, sometimes not all visible in dorsal view. Pleon seldom of more than 
two free pleonites plus pleotelson. Pleopod 1 absent in 2. One pair of 
pleopods in @, and one or two pairs of pleopods in 6 forming operculum 
over remaining respiratory pleopods. Pleopod 2 in 3 usually adapted for 
copulation. Uropods usually pedunculate, but peduncle may be reduced, 
biramous or uniramous, terminal or subterminal. 


REMARKS The suborder Asellota is usually divided into four superfamilies, 
based on pleopodal arrangement. The great majority of families, however, 


Figure 33. Virganthura crassa: A, 2; B, maxilliped; C, uropodal endopod and 
sympod; D, telson; E, uropodal exopod. 


Atlantasellus cavernicolus 15 


belong to the superfamily Janiroidea, considered to be the most advanced. In 
place of a key to the four superfamilies, the chart shown in Figure 34 illus- 
trates diagrammatically the arrangement of the pleopods in these groups. 


Superfamily Aselloidea Rafinesque-Schmaltz, 1815 


DIAGNOSIS <d: Pleopod 1, peduncles separate, rami uniarticulate; pleopod 2 
having biarticulate exopod and copulatory endopod; pleopod 3 biramous, 
opercular. 2: Pleopod 1 absent; pleopod 2 absent, or of single article; 
pleopod 3 biramous, opercular. 


Family Atlantasellidae Sket, 1979 


DIAGNOSIS’ Body resembling that of a sphaeromatid isopod. Pleon consist- 
ing of two free pleonites plus pleotelson. d: Pleopod 1 of two articles; pleopod 
2 with sympod, small exopod, and uniarticulate copulatory endopod. °: 
Pleopods | and 2 absent; pleopod 3 operculate but rami not fused. 


Atlantasellus Sket, 1979 


DIAGNOSIS. Eyes lacking. Antennule flattened, short and broad. Antenna 
elongate, peduncle of four (?five) articles, flagellum of four articles. Man- 
dibular palp of three articles; with incisor and lacinia, molar replaced by 
brushlike process. Maxillipedal palp of five articles of similar width and 
broad endite. Pereopod 1 subchelate; pereopods 2—7 ambulatory, slender, 
dactyli biunguiculate. Uropods uniarticulate, vestigial. 


Atlantasellus cavernicolus Sket, 1979 
Figure 35 


DIAGNOSIS. 6 1.1 mm. Cephalon with strong triangular rostrum, equal to 
pereonites 1-3 combined in middorsal length. Pleotelson basally broad, 
tapering to trilobed posterior margin, uropods inserted in incisions between 
median and lateral lobes. 


RECORDS Walsingham Cave, Bermuda. 


“eIOT[PSY Jopsoqgns ay} Jo sarpruueysadns inoy ay} ut ¢—] spodogyd jo uostreduiory “FE aunB1q 


\: U O ¢ GOd03 1d 
SY 
poo 


| GOd03 1d 


Gnathostenetroides pugio CE 


Superfamily Gnathostenetroidoidea Kussakin, 1967 


DIAGNOSIS 4d: Pleopod | peduncles fused, uniarticulate rami separate, op- 
ercular; pleopod 2 small, with biarticulate exopod and copulatory endopod; 
pleopod 3, broad endopod biarticulate, exopod slender, uniarticulate. 2: 
Pleopod 1 absent; pleopod 2 rami fused to form operculum; pleopod 3 as in 


OF 


Family Gnathostenetroididae Kussakin, 1967 


DIAGNOSIS Uropods with short sympod, rami relatively well developed. 3: 
Pleopod 1, protopodites short, fused, rami separate, together forming oper- 
culum covering remaining pleopods. Pleopod 2, separate, much smaller than 
pleopod 1. Pleopod 3, exopod elongate, slender; endopod uniarticulate, 
broad. 2: Pleopod 2, rami fused to form operculum covering remaining 
pleopods. 


Key to genera of Gnathostenetroididae 


1. Eyes absent. Rostrum anteriorly rounded/truncate ...... Neostenetroides 
Eyes present. Rostrum anteriorly bilobed ........... Gnathostenetroides 


Gnathostenetroides Amar, 1957 


DIAGNOSIS’ Eyes present. Rostrum well developed, anteriorly bilobed. Per- 
eonite 1, especially in d, larger than following pereopods. Pleon having one 
very short free pleonite. Pleotelson with lateral margins subparallel, ending 
in strong tooth posteriorly; posterolateral margins sinuous, rounded. Mand- 
ible in d with strong projection arising proximolateral to incisor, visible dor- 
sally; in 2, projection short, not visible dorsally. 


Gnathostenetroides pugio Hooker, 1985 
Figure 36A—C 


DIAGNOSIS 6 3.2 mm, @ 1.6 mm. Eyes of five ommatidia each. d: Per- 
eopod 1, merus with anterior margin strongly produced into narrow triangu- 
lar process; propodus broad, palm demarked by strong denticulate spine and 


78 ASELLOTA ¢ GNATHOSTENETROIDIDAE 


Figure 35. Atlantasellus cavernicolus: A, 2; B, 
pereopod |. (From Sket, 1979). 


bearing row of nine more slender spines, posterior margin setose. 2: Oper- 
culum with distal incision acute. 


RECORDS Florida Middlegrounds, Gulf of Mexico, 55 m. 


REMARKS The type of the genus, and only other species known, G. laodicense 
Amar, 1957, was recorded from the Mediterranean Sea. This species is in- 
cluded here, as several of the species first recorded from the Florida Mid- 
dlegrounds (Hooker, 1985) have since been recorded from the Caribbean. 


Neostenetroides Carpenter and Magniez, 1982 


DIAGNOSIS Pleonites 1 and 2 very short. Operculum of 2 subcircular. 
Pleopod 2 in 6 with elongate protopodite, copulatory organ prolonged by 
hyaline tonguelike structure. Pleopod 4, exopod ovate, wider than endopod. 


Neostenetroides stocki Carpenter and Magniez, 1982 
Figure 36D,E 


DIAGNOSIS 6 1.46mm, 2 1.87 mm. Eyes absent. Rostrum well developed, 
anteriorly rounded/truncate. Pleotelson wider than long, lateral margins en- 
tire, posterior margin regularly convex. Pereopod 1, propodus elongate, wid- 
ening distally, palm poorly demarked, with few spines; dactylus stout, over- 
lapping palm. Uropods unknown. 


RECORDS San Salvador, Bahamas, from Dixon Hill Lighthouse Cave. 


ASELLOTA * JANIROIDEA 79 


Figure 36. Gnathostenetroides pugio: A, 36; B, mandible; C, pereopod 1. Neostenetroides 
stocki (from Carpenter and Magniez, 1982): D, 2; E, pereopod 1. 


Superfamily Janiroidea Sars, 1899 


DIAGNOSIS <6: Pleopod 1 with elongate peduncle, occasionally fused; 
pleopod 2 with short exopod and copulatory usually elongate endopod, 
pleopods | and 2 together forming operculum; pleopod 3 endopod uniarticu- 


80 ASELLOTA * JANIROIDEA 


late, exopod biarticulate. 2: Pleopod 1 absent; pleopod 2, rami fused to form 
operculum; pleopod 3 as in 6. 


Key to families of Janiroidea 


1: Eyes (Gf present):on lateral processes of céphalon .. ..17....-. 42.025 2 
Eyes dorsolateral on cephalon, not on lateral processes ............. 6 
2. Uropods with large, easily visible sympod and rami ............... 3 
Uropods with sympod minute or absent, rami short ............... 5 


3. Pleon posteriorly markedly produced; some pereonites produced 


laterally into, fingerlike:processes: {27.2.2 Jew t ees Pleurocopidae 
Pleon posteriorly rounded, barely produced; pereonites not laterally 

produced into prominent fingerlike processes: .....-5.5.---2- ease 4 

4: \Peréopod' lSubchelate “sth 5 gas Oc as sete en ee Santiidae 

Pereopod. ] ambulatory, biunguiculate 52% os -s.6 2. .tiee ee Mexicope 


5. Pleopod 1 6 distally sagittate; anus covered by pleopod 1 d, or 


2 ber) 008 ee ee aera kta, ety Se Rr Paramunnidae 

Pleopod | 6 distally truncate; anus exposed .............. Munnidae 

6. Uropodal rami minute, smaller than squat sympod ...... Joeropsidae 

Uropodal rami elongate, sympod variable, generally elongate ....... 7 

7. Eyes lacking; pereopods all similar, ambulatory .... Microparasellidae 
Eyes usually present; pereopod | prehensile, subchelate, pereopods 

2= 7am bUlAaLOry , sch hae ven Puateeee sae ee see Re Janiridae 


Family Incertae Sedis 


Mexicope Hooker, 1985 


DIAGNOSIS_ Eyes on short lateral lobes of cephalon. Scale present on anten- 
nal peduncle. Coxae short, visible in dorsal view on pereonites 2—7. Per- 
eonites laterally acute, not markedly produced. Pereopod | ambulatory, dac- 
tylus biunguiculate. Pleon consisting of one short pleonite plus broad 
pleotelson. Uropods large, with elongate peduncle, endopod, and exopod. 


ASELLOTA * JANIRIDAE 81 


Figure 37. Mexicope kensleyi: A, 2; B, 
pereopod 1. 


REMARKS The inability to place Mexicope in a family reflects the fact that 
the arrangement of the families and genera of the Janiroidea is still unsettled. 
While having affinities with the Pleurocopidae and the Janiridae, placement 
in either of these would require redefinition of both families, making the 
Janiridae even more of a hodgepodge of phylogenetically unrelated genera. 


Mexicope kensleyi Hooker, 1985 
Figure 37 


DIAGNOSIS. 6 1.7 mm, 2 2.9 mm. Antennae slightly longer than body. Per- 
eonites and pleon laterally finely serrate. Pleotelsonic lobe between uropodal 
bases narrowly rounded, barely produced. 


RECORDS ‘Turks and Caicos Islands, 1 m; Florida Middlegrounds, Gulf of 
Mexico, 30 m. 


Family Janiridae Sars, 1899 


DIAGNOSIS Antennae longer than antennules. Mandibles with palp and 
well-developed molar. Maxilliped with articles 1—3 at least as wide as endite, 
markedly broader than articles 4 and 5. Pereopod | prehensile, subchelate, 
sexually dimorphic, larger in ¢ than in 2. Pereopods 2—7 ambulatory, dac- 
tyli biunguiculate. Coxae visible at least on three posterior pereonites. Pleon 


82 ASELLOTA * JANIRIDAE 


consisting of single free pleonite (often very narrow, short and_in- 
conspicuous) plus pleotelson. Uropods with well-developed sympod; usually 
biramous. 


Carpias Richardson, 1902 


DIAGNOSIS Cephalon with dorsolateral eyes, lacking rostrum. Coxae visible 
dorsally on pereonites 1—7. Pleon with one short pleonite lacking free lateral 
margins, plus broad pleotelson. Antennules and antennae well developed, 
latter with scale. Articles 2 and 3 of maxillipedal palp expanded. Pereopod 1 
sexually dimorphic, often relatively enormous and/or elongate in male, car- 
pochelate, carpus expanded, propodus variously armed or expanded, dac- 


Key to species of Carpias (based on pereopod 1 of mature ¢) 


L..,Propodisdistallly, Acute stoic has Rahn et Says si SS see as, seo 2 
Propodusydistally broads ooo ike oe scccs ois Sut a ona ans on 4 
2. Propodus apically acute, with proximal tooth ................ minutus 
Propodus’ lacking ‘teeth: 38 2'..18. See ane sh. IO RSE eee 3 


3. Carpus with two strong distal teeth; propodus (when chela closed) 
FEACHING. TOUMEHUS.. voles cineca sews a pe ieee eee cae oe algicola 
Carpus with two strong and one small teeth; propodus (when chela 


closed’) reaching proximallhalfof carpus 22-2: +245. 20 ec: serricaudus 
4. Propodus with distinct teeth, ots. iach yhoo es > 
Propodus lacking distinct teeth. otic cre hake ee 6 
5. Carpus distally with broadly rounded area; merus elongate-slender, 
length about four times greater than width ............. bermudensis 
Carpus lacking broadly rounded area; merus short, broader than long 
GME tho ons See t nA sd GAR socio agers Soe soe geen e: punctatus 
6. Carpus with two distal teeth; dactylus minute ................. triton 
Carpus with three distal teeth; dactylus small, but not obsolete ..... | 


7. Propodus distally truncate; carpus with middle tooth of palm distally 
faintly “bilobed 5 eS eer hee eee eee ee brachydactylus 
Propodus distally faintly bilobed; carpus with middle tooth of palm 
distally narrowly“rounded OY i2ee2 ene ee ee ae harrietae 


Carpias bermudensis 83 


tylus reduced or rudimentary, bearing two claws (biunguiculate). Pereopods 
2-7 similar, ambulatory, dactylus with three claws (triunguiculate). 
Uropods often longer than pleotelson, with relatively elongate sympod and 
rami. 


REMARKS This genus has been, and continues to be, a source of taxonomic 
problems. Several authors (e.g., Pires, 1980) have separated the species into 
the genera Carpias and Bagatus; others (e.g., Bowman and Morris, 1979) have 
synonymized them. In part, this uncertainty reflects the general uncertainty 
of the state of taxonomy in the family Janiridae. In this work, the genus 
Carpias is used to contain all the species described under the names Carpias 
and Bagatus. 

While these tiny asellotes are frequently extremely abundant in certain 
habitats (e.g., in reef-crest algal turfs; Kensley, 1983), difficulty is experi- 
enced in identifying specimens other than mature males. The first pereopod 
of the mature male is the feature best used for species separation, but varia- 
tion with maturity and geographic locality have not been investigated. With 
more detailed work, some species will undoubtedly be synonymized. 


Carpias algicola (Miller, 1941) 
Figure 38A,B 


DIAGNOSIS 6 2.9 mm, ovigerous 2 2.0 mm. Frontal margin straight. Per- 
eopod | in 6, carpus distally not much broader than proximally, with two 
teeth. Propodus reaching back to merus in mature male. Pleopod | in 4, 
rami with outer lobe distally acute but not produced. Uropod longer than 
pleotelson. Pigment in scattered red-brown chromatophores. 


RECORDS Looe Key, Florida, 1—1.5 m; Yucatan, Mexico; Carrie Bow Cay, 
Belize, 0—2 m; Puerto Rico; Jamaica; Venezuela. 
Indo-west Pacific. 


Carpias bermudensis Richardson, 1902 
Figure 38C,D 


DIAGNOSIS. 6 2.7 mm. Frontal margin straight. Pereopod | in adult ¢ al- 
most 1.6 times body length; carpus distally broadened, with rounded pos- 
terior area, palm with tooth at outer distal angle, larger tooth at midlength 
followed by deep notch; propodus with two teeth on flexor margin, widening 
to truncate distal margin. Pleopod 1 6, outer distal lobe narrowly acute, 
somewhat produced. Operculum of 2 with distal margin emarginate. 


84 ASELLOTA * JANIRIDAE 


RECORDS’ Bermuda; eastern and southern coasts of Florida, 1.5—15 m. 


Carpias brachydactylus Pires, 1982 
Figure 38E 


DIAGNOSIS <6 1.6 mm. Pereopod | 4, carpus distally between two and 
three times wider than proximal width, with strong triangular outer tooth 
defining palm, middle tooth apically faintly bifid, inner tooth rounded; pro- 
podus widening to distal truncate margin, overreaching carpal palm by short 
distance, with very low tubercle at about midlength of flexor margin. 
Pleopod | 2, outer distal lobe slightly produced, rounded. Operculum of 2 
wider than long, distal margin broadly emarginate. 


RECORDS Puerto Rico, 1.5 m. 


Carpias harrietae Pires, 1981 
Figure 38F,G 


DIAGNOSIS. 6 2.3 mm. Closely resembling C. brachydactylus. Pereopod | 6, 
carpus with posterodistal area somewhat expanded, with strong tooth defin- 
ing outer margin of palm, middle tooth rounded, inner tooth distally faintly 
bifid; propodus widening to shallowly bilobed distal margin. Pleopod | 6, 
inner distal lobe rounded, outer lobe narrowly acute. Operculum of @ with 
mediodistal margin gently concave. Uropod about twice length of pleotelson. 


RECORDS Biscayne Bay, Florida, intertidal to 2 m. 


Carpias minutus (Richardson, 1902) 
Figure 39A 


DIAGNOSIS 6 1.9 mm, ovigerous 2 1.8 mm. Pereopod | ¢, carpus distally 
widening, palm defined by strong triangular tooth, two inner teeth of palm 
separated by rounded notch; propodus with strong proximal tooth on flexor 
margin, distally produced into small triangular lobe below dactylus. Pleopod 
1 6, outer distal lobe narrowly triangular and produced well beyond inner 
lobe. Uropod subequal to pleotelson in length. 


RECORDS Bermuda, on Sargassum. 


Carpias punctatus 85 


Figure 38. Carpias algicola: A, 2; B, pereopod 1, d. Carpias bermudensis: C, D, 
pereopod 1, 3. Carpias brachydactylus: E, pereopod 1, 3. Carpias harrietae: F, G, 
pereopod 1, 6. 


Carpias punctatus (Kensley, 1984) 
Figure 39 B,C 


DIAGNOSIS 6 2.2mm, @ 2.8 mm. Dorsal integument with patchy reticulate 
pattern of large dark brown chromatophores. Frontal margin faintly convex. 


86 ASELLOTA * JANIRIDAE 


Figure 39. Carpias minutus: A, pereopod 1, 6. Carpias punctatus: B, 3; C, pereopod 
1, 3. Carpias serricaudus: D, pereopod 1, 3. Carpias triton: E, F, pereopod 1, ¢. 


Pereopod | 6, distal two-thirds parallel sided, with strong acute tooth defin- 
ing palm, and second rounded tooth; propodus with three lobe-teeth on 
flexor surface, overreaching carpus by a third of its length. Pleopod | 6, 
outer distal lobe narrowly triangular, reaching well beyond inner lobe. Oper- 
culum of 2 with distal margin shallowly concave. Uropod half length of 
pleotelson; latter with posterior margin a broadly rounded lobe between 
uropodal bases. 


RECORDS Carrie Bow Cay, Belize, intertidal to 15.2 m. 


Joeropsis 87 


Carpias serricaudus Menzies and Glynn, 1968 
Figure 39D 


DIAGNOSIS 6 1.6mm, 2 1.5 mm. Pereopod | 6, palm of carpus with two 
strong outer teeth and one short inner tooth; propodus reaching back to 
proximal half of carpus, tapering distally; dactylus obsolete. Pleopod 1 4, 
outer distal lobe acute, reaching well beyond inner lobe. Pleotelsonic margins 
very faintly serrate. Uropod about 0.7 times length of pleotelson. 


RECORDS ‘Turks and Caicos Islands, 1 m; Puerto Rico, intertidal to 1.5 m. 


Carpias triton Pires, 1982 
Figure 39E,F 


DIAGNOSIS d 2.3 mm. Very similar to C. algicola. Pereopod | d, carpus 
with two strong basally broad distal teeth; propodus extending back to merus 
in adult 6, widening to broadly rounded distal margin; dactylus minute. 
Pleopod | 6, outer distal lobe narrowly acute, reaching well beyond inner 
rounded lobe. Uropod about 1.5 times pleotelson length. 


RECORDS Carrie Bow Cay, Belize, intertidal reef crest. 


Family Joeropsidae Nordenstam, 1933 


DIAGNOSIS Cephalon free, with distinct rostrum. Molar process of mand- 
ible reduced. Maxillipedal palp articles all of similar width. Antenna short, 
peduncle dilated, flagellum reduced. Pereonites similar, wider than long. 
Pereopods similar, biunguiculate. Uropods having short squat sympod and 
very reduced rami; inserted into submedian posterior notches of pleotelson. 


Joeropsis Koehler, 1885 


DIAGNOSIS Dorsolateral eyes present. Antennule, basal article widest and 
longest, often with transparent distal dentition. Antenna, peduncular articles 
3-5 somewhat dilated, article 2 often with fringe of transparent scales; 
flagellum of about six articles, together shorter than peduncle article 5. Per- 
eonites similar, generally subequal in length and width. Pleotelson of single 
shield-shaped segment. Uropodal sympod usually with mesiodistal angle 
acute; rami reduced. 


88 ASELLOTA * JOEROPSIDAE 


Key to species of Joeropsis 


1. Lateral margins of cephalon serrate; rostrum triangular ..... personatus 
Lateral margins of cephalon entire; rostrum not triangular ......... 2 


2. Body glabrous; strong band of pigment on cephalon and pereonite 4 
5 og eR SO een eRe nen octane a Mr ak aL eS enatae te bifasciatus 


3. Rostrum evenly convex; outer uropodal ramus longer than inner 
EE eco Sch eR on eo Pee EE Ee re Noe rathbunae 
Rostrum anteriorly shallowly notched; outer uropodal ramus shorter 
tian Grime. setae. sco canes etc n Bn oie Sieve sins Weer enESeeTS coralicola 


Joeropsis bifasciatus Kensley, 1984 
Figure 40A—F 


DIAGNOSIS 6 2.5 mm, @ 2.4 mm. Body glabrous. Lateral margins of 
cephalon entire. Rostrum semicircular, with marginal flange of transparent 
teeth. Antennal flagellum of eight articles. Lateral margins of pleotelson ser- 
rate. Apex of 2 operculum blunt. Broad band of pigment on cephalon be- 
tween eyes and almost reaching posterior margin; broad band of pigment on 
pereonite 4. 


RECORDS Carrie Bow Cay, Belize, 1-6 m, often on Agaricia sp. and Porites 
sp. corals, and Halimeda sp. alga; Anguilla. 


Joeropsis coralicola Schultz and McCloskey, 1967 
Figure 40G 


DIAGNOSIS 6 2.0 mm, ovigerous 2 1.9 mm. Body setose. Lateral margins 
of cephalon entire. Rostrum anteriorly notched. Antennal flagellum of five 
articles. Lateral margins of pleotelson serrate. Apex of 2 operculum acute. 
Outer uropodal ramus shorter than inner. Pigment spread as reticulation 
over entire body. 


RECORDS Off North Carolina, on coral Oculina arbuscula; Florida Mid- 
dlegrounds, Gulf of Mexico, from sponge Agelas sp. and coral Madracis sp., 
29-33 m. 


Figure 40. Joeropsis bifasciatus: A, 6; B, pleopod 1, 3; C, uropod; D, operculum, ; 
E, antennule; F, antenna. Joeropsis coralicola: G, 3. Joeropsis personatus: H, 3. 
Joeropsis rathbunae: I, 3. 


90 ASELLOTA * MICROPARASELLIDAE 


Joeropsis personatus Kensley, 1984 
Figure 40H 


DIAGNOSIS 6 2.2 mm, @ 2.0 mm. Body glabrous. Lateral margin of 
cephalon serrate. Rostrum triangular. Lateral margins of pleotelson serrate. 
Antennal flagellum of five articles. Apex of 2 operculum acute. Outer uropo- 
dal ramus shorter than inner. Strong band of pigment on cephalon; rest of 
body with paler reticulation of pigment. 


RECORDS Carrie Bow Cay, Belize, on Porites sp. and Madracis sp. corals, 
and on Halimeda sp. alga, 1—20 m. 


Joeropsis rathbunae Richardson, 1902 
Figure 401 


DIAGNOSIS 6 1.9 mm, ovigerous 2 1.6 mm. Body setose overall. Lateral 
margins of cephalon entire. Rostrum evenly convex with flange of trans- 
parent teeth. Antennal flagellum of three articles. Lateral margins of 
pleotelson serrate. Apex of 2 operculum acute. Outer uropodal ramus longer 
than inner. Pigment in reticulation over entire body. 


RECORDS Bermuda; Florida Keys; Turks and Caicos Islands; Puerto Rico; 
Gulf of Mexico; intertidal to 36 m. 


Family Microparasellidae Karaman, 1933a 


DIAGNOSIS. Eyes lacking. Antennule much shorter than antenna. Antenna 
with scale. Pereopods all similar, with biunguiculate dactyli. Pleon of one 
free pleonite plus pleotelson. Uropods with well-developed sympod and 
rami. 


Key to genera of Microparasellidae 


1. 3, pleopod 1 narrow, not overlapping external part of pleopod 2; 
maxillipedall palp:of five articles’: 2); 34 1)--)-1- -re Microcharon 
3, pleopod 1 broad, almost completely covering pleopod 2; 
maxillipedal palp of four articles, terminal article ending in pointed 
FUEOGESS. 5c eons hats leh ole atone ee wa. electra cet eee on yet yet nee er te Angliera 


Microcharon sabulum 9] 


REMARKS All the members of this family are tiny (usually less than 2 mm), 
and most are interstitial in habit, being found in marine, brackish, and fresh- 
water environments. 


Angliera Chappuis and Delamare Deboutteville, 1955 


DIAGNOSIS Mandibular palp with two proximal articles inflated, terminal 
article, slender hooklike, articles lacking setae and spines. Maxillipedal palp 
of four articles, articles 1 and 3 elongate, article 2 short, article 4 with termi- 
nai acute process. Pleopod | in d forming broad lamella. 


Angliera psamathus Kensley, 1984 
Figure 41A—D 


DIAGNOSIS. d6 1.0mm, 2 1.0 mm. Maxillipedal endite with seven setae on 
distal margin. Posterior four pairs of pereopods with claw on dactylus dorsal 
to unguis. Uropodal endopod subequal in length to sympod. 


RECORDS Carrie Bow Cay, Belize, interstitial in intertidal sand bank. 


REMARKS _ ‘T'wo other species of Angliera have been recorded from the Carib- 
bean area: A. dubitans Stock, 1977, from Bonaire, and A. racovitzai Coineau 
and Botosaneanu, 1973, from Cuba. The reader should refer to the original 
descriptions to distinguish the species, as differences are extremely subtle. 


Microcharon Karaman, 1934 


DIAGNOSIS Mandibular palp of three articles, two distal articles bearing 
spines and/or setae. Maxillipedal palp of five articles, articles 1, 2 and 3 
expanded. d, pleopod | narrow, elongate, not obscuring pleopod 2. 


REMARKS More than 20 species of Microcharon have been described world- 
wide. The genus is unusual in that the species have been found in true ma- 
rine environments, in brackish habitats such as wells, and inland in 
freshwater. 


Microcharon sabulum Kensley, 1984 
Figure 41E—H 


DIAGNOSIS. 6 1.4mm, 2? 1.5 mm. Antennule of five articles. Inner ramus of 
maxilla 2 with pectinate spine. Pereopodal dactyli short, biunguiculate. En- 


Figure 41. Angliera psamathus: A, 3; B, right, and part of left, mandible; C, 
maxilliped; D, pleopod 1, 3. Microcharon sabulum: E, 3; F, mandible; G, 
maxilliped; H, pleopod 1, 6. 


Munna 93 


dopod of pleopod 3 with three distal plumose setae. Uropodal sympod stout, 
longer than rami. 


RECORDS Carrie Bow Cay, Belize, interstitial in intertidal sand bank. 


REMARKS ‘Two other species of Microcharon have been described from the 
Caribbean area: M. phreaticus Coineau and Botosaneanu, 1973, from intersti- 
tial freshwater on Cuba, and M. herrerai Stock, 1977, from freshwater wells on 
Bonaire. The reader should refer to the original descriptions to distinguish 
these species. 


Family Munnidae Sars, 1899 


DIAGNOSIS Body ovate. Cephalon and all pereonites free; pleon narrower 
than rest of body, longer than broad. Eyes on lateral processes of cephalon. 
Mandible with molar and incisor present; palp present or absent. Maxillipe- 
dal palp articles 2 and 3 broader than remaining articles. Pereopod | prehen- 
sile; pereopods 2—7 ambulatory. Uropods tiny, without sympod. Anus ex- 
posed, not covered by pleopods. 


REMARKS Poore (1984) has provided the most useful and recent survey of 
the genera, and especially of Munna. 


Key to genera of Munnidae 


1. Mandibular palp present; pereopod | enormous in 6, as compared 
LEE OE NTR OS Raat eee One ane ay ON ae He Munna 


Munna Kreyer, 1839 


DIAGNOSIS Body dorsally with numerous setae and/or articulating spines. 
Antennular flagellum with two distal articles each with single aesthetasc, 
terminal article minute. Mandibular molar strong, subcylindrical, distally 
truncate, with accessory setae; palp reaching beyond incisor, article 2 with 
few serrate spines. Pereopod | sexually dimorphic; pereopods 2—7 not (or 
barely) sexually dimorphic, dactyli with accessory claw. Pleopod 3, article 2 
of exopod reaching well beyond endopod. 


94 ASELLOTA * MUNNIDAE 


Munna petronastes Kensley, 1984 
Figure 42A—D 


DIAGNOSIS d 1.1 mm, 2 1.0 mm. Pereopod | in d enormously enlarged, 
carpochelate. Pleopod | in d with distolateral angles projecting, acute. Body 
with anterodorsal U-shaped pigment band, and two converging bands on 
posterior pereon. 


RECORDS Carrie Bow Cay, Belize, intertidal to 2 m, usually on corals. 


Uromunna Menzies, 1962c 


DIAGNOSIS Body with few if any dorsal setae, without articulating spines. 
Terminal antennular flagellar article not minute, bearing single aesthetasc. 
Mandibular molar strong, cylindrical, distally truncate, lacking accessory 
setae; palp present or absent. Pereopod | not sexually dimorphic, small. Per- 
eopod 2 (rarely 2—7) sexually dimorphic, carpi and propodi broader in 6 
than in 2. Pleopod 3, article 2 of exopod not reaching beyond endopod. 


Key to species of Uromunna 


1. Larger uropodal ramus parallel sided, about 3.5 times longer than 
basal width; inner uropodal ramus tiny, obscured by pleotelsonic 
TELA DORN aS. 5 oaths BP wwe eee a cee ie ie ver ke shoals har ars eet ae reynoldsi 
Larger uropodal ramus tapering, about 1.5—2.0 times longer than basal 
width; inner uropodal ramus smaller than outer, but visible beyond 


pleotelsonie mare <5 soccer ose ee ee caribea 


Uromunna caribea (Carvacho, 1977) 
Figure 42E,F 


DIAGNOSIS 6 1.5 mm, 2 1.5 mm. Propodus of pereopod 1 1.5—2 times 
longer than wide. Operculum of @ distally truncate. Shorter uropodal ramus 
visible beyond pleotelsonic margin, with single seta. Pigmentation in re- 
ticulation on cephalon and pereon; with six marginal patches on pleon. 


RECORDS ‘Turks and Caicos Islands, 1 m; Canal de la Belle Plaine, 
Guadeloupe, in water of 25%o. 


Uromunna reynoldsi 95 


Figure 42. Munna petronastes: A, 3; B, uropod; C, pereopod 1, 3; D, pereopod | ¢ 
(C and D same scale). Uromunna caribea: E, 3; F, larger uropodal ramus. Uromunna 
reynoldsi: G, pereopod 1, 6; H, larger uropodal ramus; J, 6. 


Uromunna reynoldsi Frankenberg and Menzies, 1966 
Figure 42G—I 


DIAGNOSIS <6 1.5 mm, @ 1.6 mm. Propodus of pereopod 1 two or three 
times longer than wide. Operculum of ¢ distally rounded. Shorter uropodal 
ramus obscured by pleotelsonic margin, with single seta. Pigmentation a 


96 ASELLOTA * PARAMUNNIDAE 


broad patch between eyes on cephalon, lateral bands on pereon, and anterior 
and lateral patches on pleon. 


RECORDS Sapelo Island, Georgia, in tidal saltmarsh creek; Lake 
Ponchartrain, Louisiana; Atlantic and Pacific locks of Panama Canal. 


Family Paramunnidae Vanhoffen, 1914 


DIAGNOSIS Body broad, ovate, often with laterally produced tergal or epi- 
meral plates. Cephalon recessed into pereonite 1. Eyes, if present, on lateral 
projections of cephalon. Antennule short, usually of six articles, with single 
terminal aesthetasc. Antenna never longer than body. Mandibular palp pres- 
ent or absent. Pereopod | prehensile; pereopods 2—7 ambulatory. Pleopod 1 
3 distally sagittate. Uropods with sympod minute or absent; rami tiny. Anus 
covered by pleopods. 


Munnogonium George and Stromberg, 1968 


DIAGNOSIS Eyes present on short lateral processes of cephalon. Antennal 
peduncular scale present. Coxal plates visible on pereonites 2—7. 


Munnogonium wilsoni Hooker, 1985 
Figure 43A,B 


DIAGNOSIS. <6 0.86 mm, 2 0.98 mm. Frontal margin of cephalon broadly 
rounded. Mandibular palp absent. Uropodal endopod twice length of ex- 
opod. Lateral margins of pleotelson to uropodal insertion serrate, posterior 
margin between uropodal insertions tapering to rounded apex. 


RECORDS Florida Middlegrounds, Gulf of Mexico, 55 m. 


Family Pleurocopidae Fresi and Schiecke, 1972 


DIAGNOSIS Cephalon broader than long. Eyes (or at least ocular peduncles) 
present. Mandible with or without palp; molar truncate. Maxillipedal palp 
articles narrow, less than half width of endite. At least coxae of pereonites 5— 
7 dorsally visible. Pereopods 2—7 uni- or biunguiculate. Pleopod | in 6 not 
sagittate. Uropod pedunculate, inserted laterally or slightly dorso- or ven- 
trolaterally on pleotelson; biramous, or with one ramus fused with sympod. 


Pleurocope 97 


Figure 43. Munnogonium wilsoni: A, 2; B, pereopod 1, 3. Pleurocope floridensis: C, 3; 
D, antennule 6; FE, pereopod | 6. Santia milleri: F, 3; G, pereopod 1, 3; H, 
maxilliped. 


Pleurocope Walker, 1901 


DIAGNOSIS Eyes present on lateral peduncle. Antennular peduncle of two 
articles; flagellum of four articles. Antennal peduncle of five (six) articles, 
scale lacking; flagellum of six or seven articles. Mandibular palp lacking. 
Pereopod | subchelate. Pereopods 2—7 uniunguiculate. 


98 ASELLOTA * SANTIIDAE 


Figure 44. Pleurocope floridensis: adult in dorsal 


view. 


Pleurocope floridensis Hooker, 1985 
Figure 43C—E, 44 


DIAGNOSIS. 6 1.15 mm, ¢ 0.96 mm. Body ovate, tapering posteriorly. In- 
tegument very finely tuberculate. Mesiodistal lobe on antennal peduncle ar- 
ticle 3 bearing five distal setae. Pereon lacking long dorsal setae. Pereopod 1 
subchelate, but almost carpochelate. Pleon consisting of single segment, pos- 
teriorly narrowly tapered and produced. Uropodal rami as long as sympod. 


RECORDS Turks and Caicos Islands, 1 m; Carrie Bow Cay, Belize, 3—10 m; 
Florida Middlegrounds, Gulf of Mexico, 55 m. 


Family Santiidae Wilson, 1987 


DIAGNOSIS Antennular flagellum with at most, three articles, antennular 
scale sometimes present. Pereopod 1 prehensile. Pereopods 2—7, dactyli 
biunguiculate. Coxae visible at least on pereonites 5—7. Pleon consisting of 
single short pleonite plus pleotelson. Uropods pedunculate, biramous, inser- 
ted dorsally or laterally. (One species of Santia possesses a uniramous 


uropod. ) 


Santia Sivertsen and Holthuis, 1980 


DIAGNOSIS Cephalon about twice wider than long. Eyes present. Antennu- 
lar peduncle of three articles. Pereonites laterally rounded, sometimes bear- 
ing short lateral spines. 


ASELLOTA ¢ STENETRIIDAE 99 


Santia milleri (Menzies and Glynn, 1968) 
Figure 43F—H 


DIAGNOSIS 6 and 2 1.0 mm. Eye on short lateral process of cephalon. 
Mandibular palp of three articles. Maxillipedal palp articles all of similar 
width. Pereopod | barely subchelate. Uropod with sympod well developed, 
rami prominent, well developed. 


RECORDS Carrie Bow Cay, Belize, intertidal to 30 m; Puerto Rico, 1.5 m; 
San Salvador, Bahamas, 6 m; Turks and Caicos Islands, 1 m; Anguilla; Ja- 
maica; Cozumel, Mexico; Gulf of Mexico. 

Brazil, 1-6 m. 


Superfamily Stenetrioidea Hansen, 1905a 


DIAGNOSIS’ <6: Pleopod | small, peduncles fused, rami separate, uniarticu- 
late; pleopod 2 small, copulatory; pleopod 3 biramous, opercular. 2: Pleopod 
1 absent; pleopod 2 rami and peduncle fused to form operculum; pleopod 3 


as in oC. 


Family Stenetriidae Hansen, 1905a 


DIAGNOSIS’. <d: Pereopod | frequently much bigger than in 2, with distinc- 
tive lobes and teeth. Pleopods 1 and 2 reduced; pleopod | protopodite short, 
fused, rami separate. Pleopod 2, endopod elongate, flexed, exopod short. 
Pleopod 3 exopod basally broad, distally narrowed; endopod broad, biarticu- 
late. 2: Pleopod 2, rami fused, short, covering base of pleopod 3. Uropod 
with short sympod, rami relatively well developed, styliform, of single article. 


Key to genera of Stenetriidae 


1. Rostrum narrowly triangular, spikelike; two free, very short pleonites 
anterior to pleotelson 24s eat ee ee een Stenobermuda 
Rostrum short, basally broad, anteriorly truncate or broadly rounded; 
two or three very short free pleonites anterior to pleotelson 
ee Se teh Se NCE RG, Naa eA. PUNTERS oa Stenetrium 


100 ASELLOTA * STENETRIIDAE 


Stenetrium Haswell, 1881 


DIAGNOSIS Eyes present. Cephalon broader than long. Rostrum short, 
basally broad, anteriorly truncate or rounded. Pereonites 1—4 with ante- 
rolateral projections; pereonites 5—7 projecting posteriorly. Pleotelson with 
sharp tooth anterior to small lateral notch. 


REMARKS If fresh material is not available, and color pattern is lost in pres- 
ervation, mature male material is needed as identification is based on the 
structure of male pereopod 1. 


Key to species of Stenetrium 


1. Eyes of few (not more than 10) ommatidia, not reniform ........... 2 
Eyes of many! ommmatidia, renitorm 2,,.0.103 1, boc). h . ee 3 


2. Eyes of four ommatidia; d pereopod 1 small, propodus not unusually 


21.01 (6 Mima Lea Oem eR cS gone IN MOH AE Ser RCA a minocule 
Eyes of more than four ommatidia; 3 pereopod 1, propodus broad, 

WHEL Wale soa et sian at oe Tn ee ee Nk patulipalma 

3) ‘Pleotelsonie. margins. serrate’. sess. 6 eee ee os eee 4 

Fleotelsonic*margins entire We" eh. . 4504 Be ee 5 


4. Rostrum convex, with fine marginal teeth; pereopod 1 6, propodal 


palne with*three(straight teethe Avs. 2. en iol).. atemne Sauee bowmani 
Rostrum truncate; pereopod | 3, propodal palm with two teeth, outer 

toothelongate; cued) i S20. 20a, deen). Une %, nao serratum 

5. Pereopod 1 6, carpus produced, apically acute .............. stebbingi 

Pereopod | 6, carpus produced, apically rounded ....... spathulicarpus 


Stenetrium bowmani Kensley, 1984 
Figure 45 


DIAGNOSIS 6 5.0 mm, 2 5.2 mm. Rostrum convex, with tiny marginal 
teeth. Lateral lobes of cephalon acute, margins serrate. Color pattern in 
small scattered red-brown chromatophores; irregular unpigmented patches 
on cephalon. pereonite 4, and pleon; chalky-white bands on antennae and 
uropods. d: Pereopod | propodus broad, palm with three teeth, outermost 


Stenetrium bowmani 101 


Figure 45. Stenetrium bowmani: A, 6; B, pereopod 1, 3; C, pereopod 1, 2D; 
pleopod 1, d; E, pleopod 2, 4; F, pleopod 3; G, pleopod 4; H, operculum, &. 


102 ASELLOTA ¢ STENETRIIDAE 


longest, slender. 2: Pereopod | propodus with strong denticulate spine de- 
marking palm, latter straight, with row of about seven slender spines. 


RECORDS Cozumel, Mexico; Carrie Bow Cay, Belize, 0.5—15.2 m, on algae 
and corals in reefcrest, and spur and groove zone of reef. 


Stenetrium minocule Menzies and Glynn, 1968 
Figure 46A—C 


DIAGNOSIS 6 2.8 mm, 2 3.7 mm. Eye of four ommatidia. Anterolateral 
lobes of cephalon blunt, barely produced. Rostrum poorly defined, truncate. 
3: Pereopod 1, carpus produced posterodistally into broadly rounded lobe; 
propodus broad, palm demarked by strong spine, with six low rounded teeth. 
?: Pereopod | propodus little broadened, palm demarked by strong denticu- 
late spine, bearing several more spines. 


RECORDS Puerto Rico, intertidal to 3 m; Carrie Bow Cay, Belize, intertidal 
to 36 m, from rubble, algal turfs, and seagrass. 


Stenetrium patulipalma Kensley, 1984 
Figure 46D,E 


DIAGNOSIS 6 2.0 mm, @ 2.7 mm. Eyes of about 10 ommatidia in cluster. 
Rostrum poorly defined, truncate. Two basal articles of maxillipedal palp 
not enlarged. d: Pereopod | unknown. 2: Pereopod | broadening to palm, 
and bearing row of about 12 small fringed spines. Color pattern: entire body 
with red-brown reticulation, dark transverse bars anteriorly on cephalon, 
pereonites 2 and 3, posteriorly on pereonites 4—7. 


RECORDS Carrie Bow Cay, Belize, 9.1—27.4 m; Barbados; Jamaica. 


Stenetrium serratum Hansen, 1904 
Figure 46F—H 


DIAGNOSIS. 6 4.0 mm, ovigerous 4.9 mm. Rostrum truncate. Pereonites 
1—5 with acute anterolateral angles. Pleotelsonic lateral margins with five 
teeth. d: Pereopod 1, propodus broad, palm with three teeth, outermost 
elongate, curved; dactylus reaching well beyond outermost palmar tooth. 2: 
Pereopod 1, propodus much smaller than in d, palm bearing series of about 
nine fringed spines. Color: tiny red-brown chromatophores arranged in re- 


A, 3; B, pereopod 1| 4; C, pereopod | 2. Stenetrium 
Q. Stenetrium serratum: F, 3; G, pereopod | Oe. 


Figure 46. Stenetrium minocule: 
patulipalma: D, 2; E, pereopod 1 
pereopod | Gd. 


104 ASELLOTA * STENETRIIDAE 


ticulate bands; distinctive open patches on cephalon and pereonite 1; pleon 
with two broad transverse bands. 


RECORDS Looe Key, Florida, 0.5—6 m; Turks and Caicos Islands, 1 m; 
Jamaica; Puerto Rico, intertidal to 3 m; St. Thomas, U.S. Virgin Islands; 
Carrie Bow Cay, Belize, intertidal to 15 m. 


Stenetrium spathulicarpus Kensley, 1984 
Figure 47A—C 


DIAGNOSIS. 6 4.1 mm, ¢ 4.1 mm. Rostrum truncate. d: Pereopod 1, merus 
and ischium each with setose fingerlike anterodistal projection; carpus with 
large spatulate and setose lobe almost reaching level of palm; propodal palm 
with large outer tooth and four or five low rounded teeth, broad band of setae 
near anterior margin; dactylus reaching slightly beyond palm, with band of 
setae along anterior margin. 2: Propodal palm straight, with row of slender 
spines; band of setae in similar position as in 6. Color: pigment in ill-defined 
and scattered reticulation; strong band on cephalon between eyes. 


RECORDS Carrie Bow Cay, Belize, intertidal to 36 m; Puerto Rico, 
intertidal. 


Stenetrium stebbingi Richardson, 1902 
Figure 47 D-H 


DIAGNOSIS’ 6 4.8 mm, ovigerous ¢ 4.1 mm. Rostrum truncate. Ommatidia 
of eye more bunched than in S. spathulicarpus. 6: Pereopod | variable accord- 
ing to maturity; carpus produced posterodistally into narrowly triangular, 
apically acute lobe; propodal palm poorly defined, with group of two to four 
teeth near dactylar articulation. 2: Pereopod 1, palm straight, defined by 
strong outer tooth and bearing row of six or seven low rounded tubercles. 
Color: irregular reticulation of red-brown pigment, no strong band between 
eyes. 


RECORDS’ Bermuda, 0.5—4 m; Florida Keys, 18.3 m; Bahamas, 5 m; Turks 
and Caicos Islands, 1 m; Cuba; Jamaica; U.S. Virgin Islands, 50 m; Carrie 
Bow Cay, Belize, 0.5—36 m; Gulf of Mexico. 


thulicarpus: A, 3; B, pereopod 1, 3, many setae removed; 
rium stebbingi: D, 3; E, pereopod 1, 9; F, G, H, variation 


Figure 47. Stenetrium spa 
C, pereopod 1, 2. Stenet 
in pereopod 1, d. 


106 ASELLOTA © STENETRIIDAE 


ae. 


Figure 48. Stenobermuda acutirostrata: A, 2; B, pereopod 1, °. 


Stenobermuda Schultz, 1979 


DIAGNOSIS. Eyes of few ommatidia; rostrum narrow-based, elongate and 
spikelike. Pleon consisting of two free pleonites plus pleotelson, with 
posterolateral notch marked by tooth. 


Stenobermuda acutirostrata Schultz, 1979 
Figure 48 


DIAGNOSIS 6 4.8 mm. Eyes having five ommatidia. Spikelike rostrum 
reaching well beyond anterolateral angles of cephalon. Pereopod 1, propodus 
longer than wide, palm straight, bearing eight fringed spines, posterior mar- 
gin with six spines and several setae. 


RECORDS Off Bermuda, 90 m; Turks and Caicos, | m. 


EPICARIDEA 107 


Suborder Epicaridea Latreille, 1831 


DIAGNOSIS’ Predominantly ectoparasites of marine crustaceans, feeding on 
blood. Eyes sessile, usually present in d, often reduced or lost in 2. Anten- 
nae and antennules reduced; mouthparts reduced, forming a suctorial cone 
containing pair of piercing stylets formed from modified mandibles. Maxillae 
1 and 2 reduced or lost. All mouthparts may be lost, and replaced by pro- 
boscis. d small and isopodlike. 2 undergo considerable distortion or reduc- 
tion, often to unsegmented sacs of eggs in some forms. Ostegites usually re- 
tained. Two larval mancalike stages, epicaridium and cryptoniscium (Figure 
49), characteristic of entire suborder. 


REMARKS The epicarideans are ectoparasites of other crustaceans, with the 
juveniles often using copepods as intermediate hosts. Sexual dimorphism is 
marked, the males being symmetrical with unambiguous segmentation, and 
considerably smaller than the often highly distorted females. In these, body 
segmentation is often obscured, with body segments often expanded on one 
side and reduced and compressed on the other. The marsupium of the 
ovigerous female, except in the Cryftoniscidae and Entoniscidae, is made up 
of broadly lamellar oostegites, is relatively enormous and often obscures the 
rest of the body structure. 

Crustacean hosts of the epicarideans are found in four classes: Ostracoda, 
Copepoda, Cirripedia, and Malacostraca, and in nine orders of the Mal- 
acostraca: Leptostraca, Stomatopoda, Mysidacea, Cumacea, Tanaidacea, 
Isopoda, Amphipoda, Euphausiacea, and Decapoda. The Epicaridea have 
been divided into two superfamilies, the Bopyroidea, containing families 
Bopyridae, Dajidae, and Entoniscidae, and the Cryptoniscoidea, containing 


the Crytoniscidae. 
In the Bopyridae (Figure 50), the often asymmetrical adult female shows 


some segmentation. Seven pereopods may be present only on one side, their 
number being variable on the other. This is largest of the epicaridean fam- 
ilies, containing over 400 species (Markham, 1974). Ten subfamilies have 
been recognized: the monotypic Entophilinae parasitizes galatheid crabs; six 
subfamilies, the Argeiinae, Bopyrinae, Bopyrophryxinae, Ioninae, Pseu- 
dioninae, and Orbioninae are all branchial parasites of decapod crustaceans; 
two subfamilies are abdominal parasites, the Phyllodurinae on callianassid 
mud-shrimps, and the Athelginae on hermit crabs; the Hemiarthrinae are 
known from the dorsal and ventral abdominal surfaces, and from the 
branchial chamber of caridean shrimps. 

The Dajidae are ectoparasites pelagic mysidaceans, euphausiaceans, and 
decapod caridean shrimps. Adult females are often found unattached in 
plankton and pelagic samples. When attached, dajids are found on the 


SS 
yi 


UG ae e: 
\ \ ‘SS ANY Y ASO 
x, 


¢ 


\ 


SW hs 
=a) is Soe > ae 
OP BASE ON, ey ZS 2 
WIAA ALES 


IVE 


OSS 


ON ANS 


So — 


Figure 49. A, epicaridium larva, lateral view; B, epicaridium larva, ventral view; 


C, cryptoniscium larva, ventral view (from Bonnier, 1900). 


EPICARIDEA 109 


Figure 50. A, caridean shrimp with bopyrid parasite in branchial chamber. 
Probopyrus pandalicola: B, 2 and ¢ in dorsal view, same scale; C, d enlarged; D, °, 
ventral view, eggs removed from marsupium. 


cephalothorax of the host, attached dorsally to the carapace, ventrally and 
laterally in the gill chambers and on the pereopods, or in the brood 
chambers. 

The Entoniscidae are internal parasites of decapod crustaceans, being 
found in the visceral cavity, with the parasite’s head in the position of the 
host’s gonads or hepatopancreas. Veillet (1945) demonstrated that a pore to 
the host’s branchial chamber connecting the parasite to the exterior is pres- 
ent only in hosts with mature parasites, to facilitate the release of epi- 
caridium larvae. 

The Cryptoniscidae are protandrous hermaphrodites. The female is 


110 


TABLE 2. CARIBBEAN EPICARIDEAN ISOPODS, THEIR HOSTS AND LOCALITIES 


Achelion occidentalis Hartnoll, 1966 
Microphrys bicornutus (Latreille) 


Synalpheus pectiniger Coutiére 
North Carolina to Florida; Bahamas; 
Stenorhynchus seticornis (Herbst) Hispaniola; Jamaica; Bonaire; 
Curacao; Belize; Gulf of Mexico 
Balanopleon tortuganus Markham, 1973 
Munida simplex Benedict 


Tortuga Island 


Jamaica 

Aporobopyrina anomala Markham, 1973 
Munida valida Smith 
Florida Keys; off Colombia; Gulf 


of Mexico 
Aporobopyrus curtatus (Richardson, 
1904) 
Petrochirus diogenes (Linnaeus) 
Petrolisthes armatus (Gibbes) 
Petrolisthes galathinus (Bosc) 
Petrolisthes marginatus Stimpson 
Porcellana sayana (Leach) 


Florida Keys; U.S. Virgin Islands; 


North Carolina 


Bopyrella harmopleon Bowman, 1956 
Synalpheus brevicarpus (Herrick) 
Synalpheus fritzmuelleri Coutiére 
Synalpheus hemphilli Coutiére 
Synalpheus minus Say 
Venezuela; Brazil 

Bopyrina abbreviata Richardson, 1904 
Hippolyte curacaoensis Schmitt 
Hippolyte pleuracanthus (Stimpson) 
Hippolyte zostericola (Smith) 


Argeia atlantica Markham, 1977 
Sclerocrangon jacqueti (A. Milne 
Edwards) 
Bahamas; Newfoundland 
Astalione cruciaria Markham, 1975b 
Clastotoechus vanderhorsti (Schmitt) 
U.S. Virgin Islands Bopyrione synalphei Bourdon and 
Asymmetrione clibanarii Markham, Markham, 1980 
1975d Synalpheus goodei Coutiére 
Clibanarius tricolor (Gibbes) Synalpheus bousfieldi Chace 
Florida; Bahamas; Ascension Synalpheus brevicarpus (Herrick) 
Island Synalpheus pectiniger Coutiére 
Florida; Haiti; Curacao; Gulf of 
Mexico 
Bopyrissa wolffi Markham, 1978 
Clibanarius tricolor (Gibbes) 
Clibanarius vittatus (Bosc) 


North Carolina to Florida; Belize; 
West Indies; Gulf of Mexico 
Bopyrinella thorii (Richardson, 1904) 
Thor floridanus Kingsley 
Florida; Curacao; Yucatan 
Peninsula, Mexico 


Asymmetrione desultor Markham, 1975d 
Pagurus bonairensis Schmitt 
Pagurus longicarpus Say 
Pagurus provenzanoi Forest and de 

Saint Laurent 

Bermuda; North Carolina to Florida; 
Bahamas; Puerto Rico; Gulf of 
Mexico 


Pylopagurus sp. 
North Carolina; Florida Keys; 
Curacao; Bonaire 
Azygopleon schmitti (Pearse, 1932) 
Synalpheus brooksi Coutiére 
Synalpheus hemphilli Coutiére 
Synalpheus longicarpus Coutiére 


Cabirops sp. 
Synsynella deformans Hay 
Bermuda 

Cancricepon choprae (Nierstrasz and 


Synalpheus mcclendoni Coutiére Brender a Brandis, 1925) 


Domecia acanthophora (Desbonne 
and Schramm) 
Domecia hispida Eydoux and 
Souleyet 
Eriphia gonagra (Fabricius) 
Hexapanopeus angustifrons (Benedict 
and Rathbun) 
Micropanope barbadensis Rathbun 
Neopanope packardii (Kingsley) 
Neopanope texana sayi (Smith) 
Panopeus herbstii H. Milne Edwards 
Panoplax depressa Stimpson 
Paraliomera dispar (Stimpson) 
Rithropanopeus harrisii (Gould) 
Carolinas to Florida; Bermuda; 
Curacao; Gulf of Mexico 
Cancrion carolinus Pearse and Walker, 
1939 
Panopeus herbstii H. Milne Edwards 
North Carolina; Bahamas 
Dactylokepon caribaeus Markham, 
1975c 
Iliacantha liodactyla Rathbun 
Iliacantha subglobosa Stimpson 
Dominican Republic; Costa Rica— 
Panama 
Dicropleon periclimenis Markham, 1972a 
Periclimenes americanus Kingsley 
St. Lucia Island 
Diplophryxus sp. (see Markham, 1985) 
Alpheus formosus Gibbes 
Georgia; Florida; Yucatan, Mexico 
Eophrixus subcaudalis (Hay, 1917) 
Synalpheus brooksi Coutiére 
Synalpheus goodei Coutiére 
Synalpheus hemphilli Coutiére 
Synalpheus longicarpus (Herrick) 
Synalpheus mcclendoni Coutiére 
Synalpheus pandionis Coutiére 
Synalpheus pectiniger Coutiére 
North Carolina to Florida; 
Yucatan Pensinsula, Mexico; 
Belize; Hispaniola; Curacao 


1 


Gigantione mortenseni Adkison, 1984b 
Dromidia antillensis Stimpson 
Hypoconcha sabulosa (Herbst) 
Hypoconcha spinosissima Rathbun 
Florida; Haiti; Yucatan, Mexico; 

U.S. Virgin Islands; Gulf of 
Mexico 

Hemiarthrus synalphei (Pearse, 1950) 
Synalpheus fritzmuelleri Coutiére 
Synalpheus hemphilli Coutiére 
Synalpheus longicarpus (Herrick) 
North Carolina to Florida; Haiti; 

Gulf of Mexico 

Leidya bimini Pearse, 1951 

Cyclograpsus interger (H. Milne 
Edwards) 

Pachygrapsus transversus (Gibbes) 

Sesarma ricordi H. Milne Edwards 

Bermuda; Florida Keys; Bahamas; 
U.S. Virgin Islands; Jamaica; 
Panama 

Leidya distorta (Leidy, 1855) 

Uca pugilator (Bosc) 

Uca spp. 

New Jersey to Florida; Guadeloupe; 
Trinidad 

Loki circumsaltanus Markham, 1972a 
Thor floridanus Kingsley 
Thor manningi Chace 
Southern Florida; U.S. Virgin 

Islands; Belize 
Metaphrixus carolii Nierstrasz and 
Brender a Brandis, 1931 
Hippolyte pleuracanthus Stimpson 
Southern Florida; U.S. Virgin 
Islands 

Munidion cubense Bourdon, 1972 
Munida flinti Benedict 
Munida stimpsoni A. Milne Edwards 
Cuba; Venezuela 

Munidion irritans Boone, 1927 
Munida irrasa A. Milne Edwards 
Florida Keys; Belize 


(continued ) 


112 


TABLE 2. (Continued) 


Munidion longipedis Markham, 1975a 
Munida longipes A. Milne Edwards 
Munida schroederi Chace 
East coast of Florida; Florida 

Keys; Cuba; Gulf of Mexico 

Parabopyrella lata (Nierstrasz and 

Brender a Brandis, 1929) 
Alpheus normanni Kingsley; 
Upogebia affinis (Say) 
Florida; U.S. Virgin Islands; 
Brazil 
Parabopyrella mortenseni (Nierstrasz 
and Brender a Brandis, 1929) 
Lysmata rathbunae Chace 
Lysmata wurdemanni (Gibbes) 
Florida; U.S. Virgin Islands; 
Venezuela 
Parabopyrella richardsonae (Nierstrasz 
and Brender a Brandis, 1929) 
Alpheus formosus Gibbes 
Alpheus heterochaelis (Say) 
U.S. Virgin Islands; Gulf of 
Mexico 
Parabopyrella thomasi (Nierstrasz and 
Brender a Brandis, 1929) 
Tozeuma carolinense Kingsley 
U.S. Virgin Islands 

Parapagurion imbricata Markham, 1978 
Paguristes tortugae Schmitt 
Parapagurus sp. 

Cuba; Colombia 

Parathelges foliatus Markham, 1972b 
Clibanarius vittatus (Bosc) 

Pagurus brevidactylus (Stimpson) 

Barbados; Curagao; Trinidad 

Parathelges occidentalis Markham, 

1972b 

Clibanarius tricolor (Gibbes) 

Tridopagurus sp. 

Pylopagurus corallinus (Benedict) 

North Carolina; Florida Keys; 
Bahamas; Venezuela 


Parathelges piriformis Markham, 1972b 
Paguristes oxyophthalmus Holthuis 
Pagurus brevidactylus (Stimpson) 
Pagurus provenzanoi Forest and de 

Saint Laurent 
Bermuda; Bahamas; Colombia 

Parathelges tumidipes Markham, 1972b 

Allodardanus bredini Haig and 
Provenzano 

Dardanus fucosus Biffar and 
Provenzano 

Bermuda; Jamaica 

Pleurocrypta floridana Markham, 1974 
Galathea rostrata A. Milne Edwards 
Alligator Reef, Florida 

Pleurocryptella fimbriata Markham, 

1973 
Munida constricta A. Milne Edwards 
Munida miles A. Milne Edwards 
Western Caribbean; Cuba 

Probopyria alphei (Richardson, 1900a) 
Alpheus armillatus H. Milne Edwards 
Alpheus heterochaelis Say 
Alpheus normanni Kingsley 
Alpheus viridari (Armstrong) 

North Carolina to Florida; Antilles; 
Brazil; Gulf of Mexico 

Probopyrinella latreuticola (Gissler, 1882) 
Latreutes fucorum (Fabricius) 
Bermuda; Sargasso Sea to Azores; 

North Carolina to Florida; 
Bahamas; Antilles; Gulf of 
Mexico 

Probopyrus pandalicola (Packard, 1879) 
Macrobrachium acanthurus (Wiegmann) 
Macrobrachium amazonicum (Heller) 
Macrobrachium bonelli (Nobili) 
Macrobrachium carcinus (Linnaeus) 
Macrobrachium faustinum (de Saussure) 
Macrobrachium ohione (Smith) 
Macrobrachium olfersii (Wiegmann) 
Macrobrachium surinamicum Holthuis 


Palaemon northropi (Rankin) 
Palaemon pandaliformis (Stimpson) 
Palaemonetes exilipes Stimpson 
Palaemonetes intermedius Holthuis 
Palaemonetes kadiakensis Rathbun 
Palaemonetes paludosus (Gibbes) 
Palaemonetes pugio Holthuis 
Palaemonetes vulgaris (Say) 
Periclimenes americanus (Kingsley) 
New Hampshire to Florida; 
Caribbean to Brazil; Pacific 
Panama 
Pseudasymmetrione sp. (see Adkison 
and Heard, 1978) 
Tridopagurus iris (A. Milne 
Edwards) 
Venezuela 
Pseudione affinis (Sars, 1882) 
Pandalus annulicornis Leach 
Pandalus bonnieri Caullery 
Pandalus leptorhynchus Kinahan 
Pandalus montagui Leach 
Plesionika antiguai Zariquiey 
Plesionika edwardsi (Brandt) 
Plesionika ensis (A. Milne Edwards) 
Plesionika heterocarpus (Costa) 
Plesionika martia (A. Milne 
Edwards) 
Bermuda; Northeastern Atlantic; 
South Africa; Java 
Schizobopyrina urocaridis (Richardson, 
1904) 
Periclimenes longicaudatus (Stimpson) 
Pontonia margarita Smith 
North Carolina to Florida; Belize; 
Gulf of Mexico 
Stegias clibanarii Richardson, 1904 
Clibanarius tricolor (Gibbes) 
Bermuda 


113 


Stegophryxus hyptius Thompson, 1902 
Iridopagurus sp. 
Pagurus annulipes (Stimpson) 
Pagurus bonairensis Schmitt 
Pagurus brevidactylus (Stimpson) 
Pagurus longicarpus Say 
Pagurus provenzanoi Forest and de 
Saint Laurent 
Massachusetts to Florida; Curacao 
Synalpheion giardi Coutiére, 1908 
Synalpheus longicarpus Herrick 
Yucatan, Mexico 
Synsynella choprae (Pearse, 1932) 
Synalpheus brooksi Coutiére 
Synalpheus longicarpus (Herrick) 
Synalpheus minus (Say) 
Synalpheus pandionis Coutiére 
Bermuda; North Carolina to Florida; 
Bahamas; Haiti; U.S. Virgin 
Islands; Gulf of Mexico 
Synsynella deformans Hay, 1917 
Synalpheus brooksi Coutiére 
Synalpheus longicarpus (Herrick) 
Synalpheus pectiniger Coutiére 
Bermuda; Carolinas to West Indies; 
Gulf of Mexico 
Urobopyrus processae Richardson, 
1904 
Ambidexter symmetricus Manning and 
Chace 
Processa acutirostris Nouvel and 
Holthuis 
Processa canaliculata Leach 
Processa edulis (Risso) 
Processa fimbriata Manning and Chace 
Processa tenuipes Manning and Chace 
Caribbean; Gulf of Mexico; Brazil; 
Mediterranean; Eastern Atlantic 


114 FLABELLIFERA 


reduced to a simple or lobed sac, generally without appendages. The brood- 
pouch is formed by invagination of the ventral body wall. The eggs are re- 
leased by the bursting of the sac. Cryptoniscids have been recorded as 
parasites of ostracods, cirripedes, mysidaceans, amphipods, isopods, and 
cumaceans. The majority feed on blood, but the females of some forms have 
been reported to be egg predators. 

Given the highly variable morphology of the epicarideans, and the neces- 
sity of examining large series of specimens, keys are not provided and species 
are not treated individually here. As there is a degree of genus- and species- 
specificity for the hosts, Table 2 is provided to give a clue to the possible 
identity of a specimen. The student is then advised to consult one of the 
detailed works on the group. The most useful single work on the speciose 
Bopyridae for the area covered here is Markham (1985). 


Suborder Flabellifera Sars, 1882 


DIAGNOSIS Eyes usually well developed, reduced or absent in cave forms. 
Antennules and antennae uniramous; antennal peduncle of five or six arti- 
cles. Mandible usually strong, adapted for cutting and grinding, occasionally 
for piercing; lacinia mobilis, spine-row, and molar usually present, although 
latter sometimes reduced; usually with triarticulate palp. Maxilla 1 
biramous, sometimes adapted for piercing; maxilla 2 biramous, outer ramus 


Key to families of Flabellifera 


1. Pleon consisting of four or five free pleonites plus pleotelson ........ 3 
Pleon consisting of not more than three free pleonites plus 
PlEOREISOM yee elas wie a clei maccnite. oe ork Rom eret Sines cee ete ae ee 2 


2. Pleon consisting of one or two free pleonites plus pleotelson; body 
usually dorsally strongly convex; pleopods subequal 
fae oh Ge: Ghent nara idie Met areln ome Wie aia cana te oF eet menee Sphaeromatidae 

Pleon consisting of three free pleonites plus pleotelson; body strongly 
depressed; pleopods 1—3 small, natatory, pleopods 4 and 5 large and 


broadly: ovate: jicn0 win ore | cis erie wage ac eee eee Serolidae 


3. Uropodal ramy flattened, generally not reduced” 7.2.00) - 52 4 
Uropodal rami reduced, exopod often hooklike .......... Limnoriidae 


FLABELLIFERA * AEGIDAE IPSs 


4. Pereopods 4—7 prehensile, with dactyli longer than propodi; antennae 
reduced, with no clear distinction between peduncle and flagellum 


SM as ed eS PEC PRT eS thie kala wide dan vig male subse Cymothoidae 
Pereopods 4—7 ambulatory, with dactyli shorter than propodi; antennae 
normal, peduncle and flagellum clearly distinguished ............ 5 


5. Maxilliped bearing distal recurved hooks; pereopods 1-3 strongly 


Biedeusteges Ye Pmeis aes te shies s «ts Hote oeiigadiiee ave Aegidae 
Maxilliped lacking distal recurved hooks; pereopods 1—3 ambulatory or 
AGMOSEMCAKIY, PRevensile’ ipo cf eci1 SS ayongy. ebm aise the 99 weenie « 6 


6. Maxilliped lacking, or with very reduced endite; maxilla | a strongly 
RetNeABe MENA D. Sei Segoe Ai an eter ae wists aos Sys Corallanidae 
Maxilliped with strong endite; maxilla 1 not strongly falcate ........ | 


7. Mandibular incisor distally narrowed, lacinia lacking; maxilla 1 slender 

and elongate, with 3—5 distal hooked spines ........ Tridentellidae 
Mandibular incisor distally broad, cusped; maxilla | relatively broad, 

with several distal spines and setae .................. Cirolanidae 


usually consisting of two lobes. Pereopods generally ambulatory, sometimes 
prehensile; pereopods 1 and 2 subchelate only in Serolidae, ancinine 
Sphaeromatidae, and some Cirolanidae; posterior pereopods sometimes sec- 
ondarily natatory in some cirolanids. Pleon consisting of as many as five free 
pleonites plus pleotelson, but pleonites variously fused in several families. 
Five pairs of pleopods usually present. Uropods lateral, usually forming tail- 
fan with pleotelson. 


REMARKS This suborder contains a large group of diverse families, largely 
held together by primitive features such as the tailfan structure. Future work 
will undoubtedly show the Flabellifera to be an artificial polyphyletic group. 


Family Aegidae Leach, 1815 


DIAGNOSIS Dorsal integument usually unornamented. Coxae distinct on 
pereonites 2—7. Eyes usually present, large, often almost, or complete con- 
tiguous. Mandible lacking lacinia mobilis, spine-row, and molar. Maxilla | 
slender, with apical spines. Maxilla 2 with two terminal unequal lobes bear- 
ing apical spines. Maxillipedal palp of two, three, or five articles. Pereopods 
1—3 prehensile, with dactyli strongly curved; pereopods 4—7 ambulatory. 


\ 


116 FLABELLIFERA * AEGIDAE 


Pleopods biramous, bearing plumose marginal setae. Uropods forming tail- 
fan with pleotelson. Pleon of four or five free pleonites plus pleotelson. 


REMARKS Although these large isopods (up to 60 mm) are often referred to 
as fish parasites, Brusca (1983) prefers the term “carnivorous scavengers and 
micropredators,” as they attach to fish hosts infrequently and only long 
enough to feed. When feeding, they engorge themselves on the host’s blood. 
Aegids show almost no host- (or rather prey-) specificity, being opportunistic 
feeders, and are most frequently captured by bottom trawls on the ocean bed. 
In ovigerous females, the maxillipedal articles become expanded and, along 
with the anterior oostegites, cover the buccal field, thereby making feeding 
impossible. 


Key to genera and subgenera of Aegidae 


1. Maxillipedal palp of two or three articles; frontal lamina small, narrow 
SABO Aes Use bs season, CR ae ee eae cs ase Rocinela 


2. Antennular peduncle articles 1 and 2 expanded; cephalon lacking true 
rostrum, not completely separating antennular bases .... Aega (Aega) 
Antennular peduncle articles 1 and 2 not expanded; cephalon with true 
rostrum completely separating antennular bases ... Aega (Rhamphion) 


Aega Leach, 1815 


DIAGNOSIS Eyes large, contiguous or separate. Cephalon with or without 
true rostrum. Frontal lamina broad, separating bases of antennae. Mandibu- 
lar palp article 2 elongate. Maxilla 1 bearing strong apical and subapical 
spines. Maxilla 2 of two usually unequal lobes bearing stout spines. Max- 
illipedal palp of four or five articles, terminal article often small, with setae or 
recurved spines; article 4 with stout recurved spines; endite small, seldom 
reaching beyond palp article 2. Pleon not much narrower than pereon. 


REMARKS Brusca (1983) published a useful account of the genus Aega in the 
Eastern Pacific. 


Aega (Aega) ecarinata Ie Wg 


Key to species of Aega (Aega) 


RP es eONP EI UNOS Sete ics aa. 0 aio ls isbn ua YY Gig ai Ba tepore en ok 2 deshaysiana 
BERS SSCA ALE Be eH Oe ah be Sa oo oe ops ayo ey herve divine ole bon eR ecarinata 


Key to species of Aega (Rhamphion) 


1. Posterior margin of pleotelson distinctly dentate .............. dentata 
Posterior margin of pleotelson at most faintly crenulate ....... tenuipes 


Aega (Aega) deshaysiana (H. Milne Edwards, 1840) 
Figure 51A 


DIAGNOSIS ? 18.0 mm. Eyes large, contiguous. Cephalon with frontal mar- 
gin acute to subacute. Frontal lamina large, shield shaped. Antennular ped- 
uncle articles 1 and 2 not expanded; flagellum of more than 15 articles. 
Uropodal endopod with deep notch in lateral (outer) margin. Pleotelson with 
basal width subequal to middorsal length, triangular, lateral margins faintly 
to markedly convex, tapering to narrowly rounded to subacute apex. 


RECORDS Cuba; Yucatan Peninsula, Mexico; Gulf of Mexico. 

Azores; Cape Verde Islands; Tristan da Cunha; Mediterranean; St. Paul and 
Amsterdam Islands; Seychelles; east coast of South Africa; Philippines; Ja- 
pan; Hawaii; northeast Australia; Tasmania; Cocos Islands; Costa Rica. 


REMARKS This species is more familiarly known in the Caribbean region as 
Aega antillensis Schioedte and Meinert, 1879. 


Aega (Aega) ecarinata Richardson, 1898 
Figure 51B 


DIAGNOSIS ¢ 21.0 mm. Eyes well separated. Articles 1 and 2 of antennular 
peduncle expanded. Propodus of pereopod 3 with posterodistal lobe. Uropo- 


118 FLABELLIFERA * AEGIDAE 


Figure 51. A, Aega (Aega) deshayesiana; B, Aega (Aega) ecarinata; C, Aega (Rhamphion) 
dentata; D, Aega (Rhamphion) tenuipes. 


dal exopod narrower than endopod; latter distally truncate, lacking marginal 
notch. Pleotelson dorsally smooth, with posterior margin broadly trilobed. 


RECORDS Bahamas, 776 m; Puerto Rico; Gulf of Mexico, 176 m. 


Rocinela 119 


Aega (Rhamphion) dentata Schioedte and Meinert, 1879 
Figure 51C 


DIAGNOSIS @ 7.5 mm. Eyes large, just contiguous in midline. Frontal lam- 
ina distally acute. Uropodal exopod shorter than and half width of endopod; 
latter with lateral margin entire. Pleotelson with two obscure dorsal depres- 
sions anteriorly; posterior margin crenulate with seven teeth. 


RECORDS Cuba. 


Aega (Rhamphion) tenuipes Schioedte and Meinert, 1879 
Figure 51D 


DIAGNOSIS 2 11.5 mm. Eyes large, contiguous. Frontal lamina distally 
broadly rounded. Uropodal exopod shorter and narrower than endopod; lat- 
ter with entire lateral margin. Pleotelson dorsally smooth; posterior margin 
evenly and broadly convex, obscurely crenulate. 


RECORDS Cuba. 


Rocinela Leach, 1818 


DIAGNOSIS Cephalon with short rostrum sometimes covering antennular 
bases. Eyes well developed. Frontal lamina small, often indistinct. Mandibu- 
lar palp of three articles, article 1 elongate. Maxillipedal palp of two or three 
articles. Pereopods 1—3 usually with spine-bearing expanded lobe on pos- 
terior margin of propodi. 


Key to species of Rocinela 


EV CSMEOILIC MOUS 57.07724. sc Gli DARSs SOE UL ie ae ieaies ten ee ere oculata 
By CS et COMM GUOUS. % 5. 3d. ocie sueltns sfliogy a brecee Sle Gisiaacs & ccehans ss OR ere: 2 
2. Cephalon produced anteriorly into broadly rounded rostrum. . cubensis 
Cephalon lacking obvious broadly rounded rostrum ............... 3 
3. Eyes well separated, cephalon anteriorly broadly triangular .... signata 


Eyes barely separate, cephalon anteriorly narrowly triangular —_ insularis 


120 FLABELLIFERA * AEGIDAE 


Rocinela cubensis Richardson, 1898 
Figure 52A 


DIAGNOSIS. 6 16 mm. Cephalon with two small tubercles between well- 
separated eyes; rostrum broadly rounded, extending anteriorly, very ob- 
vious. Flagellum of antenna with about 15 articles. Propodi of pereopods 1—3 
with two spines. Pleotelson basally wider than middorsal length, lateral mar- 
gins convex, apex rounded. 


RECORDS Off Cuba, 290 m. 


Rocinela insularis Schioedte and Meinert, 1879 
Figure 52B 


DIAGNOSIS Ovigerous 2 24.5 mm. Eyes medially barely separated but not 
contiguous. Flagellum of antenna of more than 12 articles. Propodus of per- 
eopods 1-3 with two to four spines on posterior lobe. Uropodal endopod 
barely reaching beyond pleotelsonic apex. Pleotelson basally slightly wider 
than middorsal length, lateral margins convex, apex rounded. 


RECORDS Florida Keys; West Indies; between Mississippi delta and west 
coast of Florida, Gulf of Mexico; 550 m. 


Rocinela oculata Harger, 1883 
Figure 52C 


DIAGNOSIS ¢ 21.0 mm. Eyes contiguous. Cephalon with rostrum truncate 
in dorsal view. Antennal flagellum with 12 articles. Propodi of pereopods 1—3 
with six to eight spines on lobed posterior margin. Pleotelson with basal 
width subequal to middorsal length; posterior margin broadly rounded. 


RECORDS Off Georgia; Gulf Stream off Florida, 360—400 m; Puerto Rico, 
84 m; Gulf of Mexico, 380-750 m. 
New South Wales, Queensland, Australia, 450—630 m. 


Rocinela signata Schioedte and Meinert, 1879 
Figure 52D 


DIAGNOSIS Ovigerous 2 13.0—15.0 mm. Cephalon anteriorly broadly tri- 
angular, produced over bases of antennules. Eyes widely separate. Flagellum 
of antenna with 10 or 11 articles. Pereopods 1—3, propodi unarmed or with 


Figure 52. A, Rocinela cubensis; B, Rocinela insularis; C, Rocinela oculata; D, Rocinela 
signata. 


122 FLABELLIFERA * CIROLANIDAE 


single spine on posterior margin. Pleotelson with posterior margin evenly 
and broadly rounded; usually with inverted W-shaped band of pigment. 


RECORDS Florida Keys, shallow infratidal—4 m; Tortugas, from gills of jew- 
fish Epinephelus itajara, mutton snapper Lutjanus analis; U.S. Virgin Islands, 
on mutton snapper Lutjanus analis, on yellowfin grouper Mycteroperca venenosa; 
Bahamas, on sheepshead Archosargus probatocephalus, on mutton snapper Lut- 
janus analis, on blackfin snapper Lutjanus buccanella, on queen triggerfish Bal- 
istes vetula, on saucereye porgy Calamus calamus; Jamaica, on French grunt 
Haemulon flavolineatum, hogfish Lachnolaimus maximus, on parrotfish Sparisoma 
viride; Haiti; Yucatan Peninsula, Mexico, 60—93 m, on gills of tiger shark 
Galeocerdo cuvieri; Puerto Rico, in gill slits of southern stingray Dasyatis amer- 
icana, in gill slits of nurse shark Ginglymostoma cirratum; Jamaica; Carrie Bow 
Cay and Blue Ground Range, Belize, 0.5—2 m, on jolthead porgy Calamus 
bajonado and sheepshead porgy Calamus penna, on peacock flounder Bothus lu- 
natus, on queen triggerfish Balistes vetula, on Caranx sp., on barracuda 
Sphyraena barracuda, on hogfish Lachnolaimus maximus, on mutton snapper Lut- 
janus analis; Venezuela, on Orthopristis ruber, on Haemulon steindachneri; Sur- 
inam, on gills of sheepshead porgy Calamus penna; Gulf of Mexico off Florida, 
shallow infratidal—55 m, on red grouper Epinephelus morio, on Lutjanus black- 
fordi, on black grouper Mycteroperca bonaci, on clearnose skate Raja eglanteria. 

Pacific records: Southern California and Gulf of California; Socorro Is- 
land; Panama; Costa Rica. 


REMARKS While often taken from fish hosts (sometimes in the gill cham- 
ber), this species is equally frequently found freeliving in shallow water over 
sand and coral rubble. The species will also attach itself to humans, inflicting 
a sharp bite as it tries to feed. 


Family Cirolanidae Dana, 1852 


DIAGNOSIS Eyes when present, relatively small, lateral. Frontal lamina 
present. Mandible with tridentate incisor, lacinia mobilis, blade- or sawlike 
molar, and palp. Maxillipedal palp of five articles, endite present. Coxal 
plates present on pereonites 2—7, distinctly separated by suture from tergite. 
Pereopods generally ambulatory, although anterior three pairs prehensile in 
some genera, and posterior four pairs natatory in some genera. Pleon of five 
free pleonites plus pleotelson in most genera; pleonite 5 with free lateral mar- 
gins or overlapped by pleonite 4. Pleopods membranous, lacking ridges or 
folds. Uropods situated at anterolateral angles of pleotelson, freely articulat- 
ing, both rami well developed, mobile. 


FLABELLIFERA * CIROLANIDAE 123 


REMARKS Of the many recent publications on the cirolanids, the most com- 
prehensive is that of Bruce (1986) on the cirolanids of Australia. Botosa- 
neanu, Bruce, and Notenboom (1986) tabulate all the known troglobitic 
cirolanids of the world. 


Key to subfamilies of Cirolanidae 


1. Clypeus projecting; pleonite 5 with free lateral margins (except in 
PROP ITIP) ML ie eee eae Ws ARE, Spm 4 le UO wre «TS Eurydicinae 
Clypeus flattened, not projecting; pleonite 5 lacking free lateral margin, 
avenapped*by-pleonite’ 4) 2020 oe Pek. ek el dav Se ES 2 


2. Pereopods 1—3 with ischium and merus not anterodistally produced; 
antennal peduncular articles 4 and 5 subequal; secondary unguis 
present on percopodal dactyli- y+. -Pe Ts eee ee ae. Cirolaninae 

Pereopods 1—3 with ischium and merus anterodistally produced; 
antennal peducular articles 3 and 4 subequal; no secondary unguis 
GHePEneOPOUAlGAClYIN ' 8 s1s.8 sete Picea sod Mas oe eo eA a a ares Conilerinae 


Subfamily Cirolaninae Dana, 1852 


DIAGNOSIS Frontal lamina short, flat. Clypeus flattened, not projecting. 
Antennal peduncular articles 4 and 5 subequal, longer than articles 1—3. 
Pereopods with secondary unguis on dactyli. Penes reduced or absent. 
Pleonite 5 always overlapped by pleonite 4. Pleopod 2 in d with copulatory 
stylet articulating basally. 


Key to genera of Cirolaninae 


1. Pleopods having accessory branchial filaments ............ Bathynomus 
Pleopods lacking accessory branchial filaments .................... 2 
eet COO ene LCUMNONMIGE 0.8 4 hon edicles averse Mele d gkn ek Aew sine Os es 7 
Picopad -ltnereperculiforni shoe! 220m. Gis he eh. Bi gee oreei . 3 
3. Pleopods 3-5, endopods lacking, or with very few, marginal setae ... 4 
Only endopod of pleopod 5 lacking marginal setae ........... Cirolana 


(continued ) 


124 FLABELLIFERA * CIROLANIDAE 


Key to genera of Cirolaninae (Continued) 


4. Merus of pereopod | posterodistally produced; merus of pereopods 2 


and 3 anterodistally produced s'.. c 2 sn: eae meee Bahalana 
Meri ‘of pereopods. 1—3- not markedly produced ......4..5022 +5598 5 

5. ‘Animal able .to.conglobates 2 cepts pene Yee ee tae Creaseriella 
Animal unabletejconclobatelusescis tastes alowed: Asean 6 

6G. Mandibular palp directed’ antenorly” ... 2. 5..5.-.-----.9:- Anopsilana 
Mandibular palp-directed posteriorly .....-5..-24-.200-5458 Haptolana 

7. Pleopod 1, exopod longer and broader than endopod ..... Oncilorpheus 
Pleopod 1, endopod longer and broader than exopod ...... Calyptolana 


Anopsilana Paulian and Delamare Deboutteville, 1956 


DIAGNOSIS Body unable to conglobate. Eyes present or absent. Frontal 
lamina well developed, as long as broad, or longer than broad, anteriorly 


Key to species of Anopsilana 


1. Estuarine-brackish water species; integument pigmented when alive 2 


Cave species;lacking antegumental: pigment, |< f<.0 6 f.6, we2ho cee etm ory 3 
2. Frontal lamina distally rounded; projectune {25.°..-. 25. 5-22. browni 
Frontal lamina distally acute, not projecting .................. jonest 
3. Posterior margin of pleotelson with 10 or more spines .............. 4 
Posterior margin of pleotelson with less than 10 spines ............. 5 


4. Posterior margin of pleotelson with 10 spines; found in cave on Cuba 
cubensis 


Posterior margin of pleotelson with 10—12 spines; found in cave on 
Haiti 


©) ehce: eB, 6s! epee (a, 8/8) 6. 6) (aL 6) Sm, .0) «0! ee) ae |e) iWiqiej) © fefle| (elim) se) 1) 8 le a) e's) a) .e) (ell eie/ce! e-em imam 


acanthura 


5. Posterior margin of pleotelson with eight spines; found in cave on 
Grand Caymai: :.6 aka 5 tase pyedonsthe aioeie aie ee eee crenata 
Posterior margin of pleotelson with four spines; found in cave on Haiti 
radicicola 


Bie 68) 0) Ge Om ee Oe) Sh Si ep ee, wel in esfee) ee) a6) a) fo) @ (6) eo (ame. le louie. ote 6) (a) ese mies (ee cee ane 


Anopsilana crenata 125 


somewhat expanded. Antennular peduncle of two articles. Maxillipedal en- 
dite with two coupling hooks. Pereopod | prehensile, pereopods 2—3 weakly 
prehensile, pereopods 4—7 ambulatory. Pleopod 2 d, copulatory stylet artic- 
ulating at base of endopod. Pleopods 3-5, exopods biarticulate, endopods 
lacking marginal setae. Uropodal sympod produced along mesial margin of 
endopod. 


Anopsilana acanthura (Notenboom, 1981) (was fla L Lome. 
Figure 53A,B 


DIAGNOSIS 6 7.0 mm. Lacking eyes and integumental pigment. Frontal 
lamina anteriorly rounded. Posterior margin of pleotelson with 10—12 spines. 


RECORDS Well at Marigot, Haiti. 


Anopsilana browni (Van Name, 1936) 
Figure 53C,D 


DIAGNOSIS. 6 11.1 mm, 2 10.0 mm. Eyes well developed, pigmented. Dor- 
sal integument strongly pigmented with red-brown chromatophores. Frontal 
lamina as wide as long, anteriorly rounded. Cephalon with two fused mid- 
dorsal tubercles near posterior margin. Pereonites and pleonites each with 
row of tubercles near posterior margin. Pleotelson triangular, with scattered 
dorsal tubercles, apex rounded, with eight spines. 


RECORDS River in Santa Clara Province, Cuba (freshwater); Sittee River 
and Salt Creek, Stann Creek District, Belize (brackish water). 
Golfo de Nicoya, Pacific Costa Rica, in red mangroves. 


Anopsilana crenata Bowman and Franz, 1982 
Figure 53E,F 


DIAGNOSIS 6 6.2 mm. Lacking eyes and integumental pigmentation. 
Frontal lamina longer than wide, anteriorly rounded. Posterior margin of 
pleotelson with eight spines. 


RECORDS West Bay Cave, Grand Cayman Island. 


126 FLABELLIFERA * CIROLANIDAE 


Figure 53. Anopsilana acanthura: A, 2; B, anterior cephalon. Anopsilana browni: C, &; 
D, anterior cephalon. Anopsilana crenata: E, 2; F, anterior cephalon. Anopsilana 
cubensis: G, 2; H, anterior cephalon. 


Anopsilana cubensis (Hay, 1903) 
Figure 53G,H 


DIAGNOSIS ¢ 7.0 mm. Lacking eyes and integumental pigmentation. 
Frontal lamina longer than wide, anteriorly expanded, rounded. Posterior 
margin of pleotelson with 10 spines. 


Bahalana 127 


RECORDS Caves in provinces of Pinar del Rio, La Habana, Matanzas, and 
on Isla de Pinos, Cuba. 


Anopsilana jonesi Kensley, 1987 
Figure 54A—D 


DIAGNOSIS 6 7.2 mm, ovigerous ¢ 5.9 mm. Eyes well developed and pig- 
mented. Dorsal integument strongly pigmented with almost solid central 
area on pereonites 1—7. ¢ cephalon with three low tubercles near posterior 
margin; pereonite 1 with four to six low tubercles. 2 lacking tubercles on 
cephalon or pereonite 1. Pereonites 2—7 with low submedian longitudinal 
ridges near posterior margin. Frontal lamina narrow, pentagonal, anteriorly 
acute, not projecting. Posterior margin of pleotelson with 9 or 10 spines. 


RECORDS Salt Creek, and Sittee River, Stann Creek District, Belize, in es- 
tuarine mangroves. 


Anopsilana radicicola (Notenboom, 1981) 
Figure 54E,F 


DIAGNOSIS. 6 6.3mm, 2 6.5 mm. Lacking eyes and integumental pigmen- 
tation. Frontal lamina longer than wide, anteriorly expanded and rounded. 
Posterior margin of pleotelson with four spines. 


RECORDS Source Débarasse, a spring near Jérémie, Haiti. 


Bahalana Carpenter, 1981 


DIAGNOSIS Eyes lacking. Frontal lamina basally triangular, anteriorly nar- 
rowed into poorly developed carina. Pereopods 1—3 prehensile, dactyli and 
propodi relatively elongate; pereopod 1 carpus small, almost concealed; 
merus with strong posterodistal extension almost reaching dactylar base and 
armed with spines. Pereopods 2 and 3, meri with elongate anterodistal exten- 
sion, meri and carpi with shorter posterodistal extensions. Pereopods 4—7 
slender, ambulatory. Pleopod 2 in d with copulatory stylet articulating 
basally on endopod. Pleopods 3—5, exopods biarticulate, endopods with few 
distal marginal setae, or lacking setae. Pleonite 5 with free lateral margin, 
hardly overlapped by pleonite 4. 


128 FLABELLIFERA ¢ CIROLANIDAE 


Key to species of Bahalana 


1. Pleopods 3—4, endopods lacking setae; maxillipedal endite with one 
coupling hook sat tin Ge SOMERS als haat on tes eae eae sete cardiopus 
Pleopods 3—4, endopods with few setae; maxillipedal endite with one or 
twovcoupling NOOKS 4.....025 eae. s: aeele «Burg te eo eee 2 


2. Antennular peduncle, article 3 longest; maxillipedal endite with two 
CouplingyhoOoksSgee 2 ~ seica sucht a age = «bac. ae ee geracei 

Antennular peduncle, article 2 longest; maxillipedal endite with one 
COUPER HOON teste san css ask, cana tale, Be 6 bt we os en mayana 


Bahalana cardiopus Notenboom, 1981 
Figure 55A,B 


DIAGNOSIS ¢ 10.0 mm. Maxillipedal endite with single coupling hook. Per- 
eopod 1, meral projection bearing five distal spines. Pleopods 3—5, endopods 
lacking marginal setae. Uropodal exopod half width of endopod, four spines 


on outer margin. 


RECORDS Mount Misery Cave, Mayaguana Island, Bahamas. 


Bahalana geracei Carpenter, 1981 
Figure 55C—G 


DIAGNOSIS 15.0 mm, d 8 mm. Maxillipedal endite with two coupling 
hooks. Pereopod 1, meral projection bearing seven distal spines. Pleopods 3— 
5, endopods with 9-13 distal marginal setae; pleopod 5 endopod with four 
distal setae. Uropodal endopod bearing few spines at outer distal margin, 
margin not serrate, lacking distinct apex; exopod half width of endopod, with 
four spines on outer margin. 


RECORDS Lighthouse Cave, San Salvador Island, Bahamas. 


Bahalana mayana Bowman, 1987 
Figure 55H 


DIAGNOSIS 2 8.4 mm, d 10.0 mm. Clypeus acutely pointed. Antennular 
peduncle, article 2 longest. Maxillipedal endite with one coupling hook. Per- 


Bathynomus 129 


B 


Figure 54. Anopsilana jonesi: A, 36; B, anterior cephalon; C, pleopod 2, 3; D, 
pleopod 3. Anopsilana radicicola: E, 3; F, anterior cephalon. 


eopod 1, meral projection rudimentary. Pleopods 3—4, endopods with few 
marginal setae, pleopod 5 lacking marginal setae. Uropodal exopod narrow, 
73 width of endopod; endopod with distal margin slightly concave. 


RECORDS Anchialine caves on Cozumel Island and at Tulum, Yucatan 
Peninsula, Mexico. 


Bathynomus A. Milne Edwards, 1879 


DIAGNOSIS Frontal lamina triangular; clypeus projecting anteriorly. Anten- 
nal peduncle with articles 3 and 4 subequal, article 5 longest. Maxillipedal 
endite with four to seven coupling hooks. Pereopods 1—3 with anterodistal 
margins of ischia and meri produced. Pleopods with all rami bearing margi- 
nal setae; endopods bearing accessory gills at bases. Posterior margin of 
pleotelson dentate. 


Figure 55. Bahalana cardiopus: A, 2; B, pleopod 3. Bahalana geracei: C, 2; D, 
pleopod 3; E, pereopod 1; F, pereopod 2; G, anterior cephalon. Bahalana mayana: 
Hy 2. 


Calyptolana 131 


Figure 56. Bathynomus giganteus. 


Bathynomus giganteus A. Milne Edwards, 1879 
Figure 56 


DIAGNOSIS Up to 280 mm. Large pigmented eyes present, not visible dor- 
sally. Antennule with small exopod distally on peduncular article 3. Pleopods 
with marginal setae on all rami; pleonites 3 and 4 with epimera produced 
posteriorly. Posterior margin of pleotelson with median tooth and five or six 
teeth on each side. 


RECORDS Gulf of Mexico; Caribbean; Bahamas; Florida Keys; 360—2300 


m. 


REMARKS Gut-content analysis of these deep-water giants has shown them 
to be scavengers, commonly feeding on dead fish, cephalopods, crabs, and 
polychaete worms. 


Calyptolana Bruce, 1985 


DIAGNOSIS. All pereopods ambulatory; each dactylus with secondary un- 
guis. Pleopod | operculate, longer than following pleopods. All pleopodal 
rami except endopod of pleopod 5 with marginal setae. 


132 FLABELLIFERA ¢ CIROLANIDAE 


Calyptolana hancocki Bruce, 1985 
Figure 57 


DIAGNOSIS 3.0 mm. Body dorsally strongly convex. Cephalon with small 
rostrum curving ventrally to meet frontal lamina; latter pentagonal. Eyes 
small, well pigmented. Coxae of pereonites barely produced. Pleonite 5 lack- 
ing free lateral margins. Pleotelson with broadly rounded posterior margin. 
Uropodal sympod produced along mesial margin of endopod; exopod 
slightly less than half length of endopod; latter distally rounded. 


RECORDS Dominican Republic; Aruba Island, Netherlands Antilles, 43.2 


m. 


Cirolana Leach, 1818 


DIAGNOSIS’ Frontal lamina usually twice as long as wide, not projecting; 
clypeus flat. Mandible with strong incisor, dentate molar, palp of three arti- 
cles. Pereopods all ambulatory. Pleon consisting of five free pleonites plus 
pleotelson, pleonite 5 overlapped laterally by pleonite 4. All pleopodal rami 


Key to species of Cirolana 


1. Pleotelson posteriorly very broad to subtruncate .................. 2 
Pleotelsom postenionly narmowed Hills, 29g Ie 2 ase eee 3 See ee 4 


2. Uropodal endopod broad, distally rounded, lacking marginal spines 
aoa dp hs SRhe nas Spe anc spovena ve wy oe a NRL ann a eee a eee ca obtruncata 


earn elie 20h OE Ta aaa One GARE Chg) UR As Nich tr eee 3 


3. Posterior margin of pleotelson bearing spines; uropodal endopod not 
lenibate eh Fig a. Tea BT TAI ee Le ee ke minuta 
Posterior margin of pleotelson faintly crenulate, lacking spines; 
uropodal ‘endoped strongly dentate 52. 5.-2..5. =1 nen crenulitelson 


4. Uropodal endopod evenly tapering to acute apex; uropodal exopod 
length about four times greatest width. 2%. 425... 6.26427 e05 albidoida 
Uropodal endopod, outer margin convex; uropodal exopod length less 
thanfour times 'greatest width yw ieHeliin Se eee parva 


Cirolana crenulitelson 133 


Figure 57. Calyptolana hancocki: A, 2; B, ventral pleon. 


bearing marginal setae except endopod of pleopod 5. Copulatory stylet on 
endopod of ¢ pleopod 2 inserted proximally. 


Cirolana albidoida Kensley and Schotte, 1987 
Figure 58A—C 


DIAGNOSIS. <6 7.8 mm. Integument sparsely pitted. Antenna reaching pos- 
teriorly to pereonite 3. Uropodal endopod triangular, evenly tapering, mar- 
gins serrate; exopod length about four times greatest width, apically acute. 
Pleopod | 3d, endopod markedly narrowed in distal half. Pleopod 2 d, copu- 
latory stylet reaching beyond rami by about half its length. Pleotelson with 
sides gently convex, tapering to rounded posterior margin bearing eight 
spines and three or four small serrations anterior to first spine. 


RECORDS Off Lucaya, Grand Bahama, 180-220 m. 


Cirolana crenulitelson Kensley and Schotte, 1987 
Figure 58D-F 


DIAGNOSIS <6 7.0mm, 2 7.0 mm. Antenna reaching posteriorly to anterior 
of pereonite 3. Pleopod 2 6, copulatory stylet reaching by '/s its length be- 


134 FLABELLIFERA * CIROLANIDAE 


Figure 58. Cirolana albidoida: A, 3; B, uropod; C, pleotelsonic apex. Cirolana 
crenulitelson: D, 2; E, pleotelsonic apex; F, uropod. Cirolana minuta: G, 3; H, 
anterior cephalon; /, uropod. 


yond rami. Uropodal endopod with mesial margin broadly convex, serrate, 
apically acute; exopod about 2.5 times longer than wide, mesial margin 
weakly convex, apically acute. Posterior margin of pleotelson truncate, 
faintly crenulate, lacking spines. 


RECORDS Carrie Bow Cay, Belize, 36 m. 


Cirolana parva 135 


Cirolana minuta Hansen, 1890 
Figure 58G-—I 


DIAGNOSIS 6 8.9 mm, ¢ 5.0 mm. Antenna reaching posteriorly to per- 
eonite 3. Pleopod 2 d, copulatory stylet reaching by about '/ of its length 
beyond rami. Uropodal endopod, mesial margin broadly convex, with apical 
angle of about 90°; exopod about three times longer than wide, parallel sided 
for proximal two-thirds, with marginal spines, apically narrowly rounded. 
Pleotelson with posterior margin subtruncate to broadly rounded, with about 
eight marginal spines. 


RECORDS St. Thomas, U.S. Virgin Islands; Tobago; 180-220 m. 


Cirolana obtruncata Richardson, 1901 
Figure 59A,B 


DIAGNOSIS. 6 11.3 mm, 2 11.0 mm. Antenna reaching posteriorly to mid- 
dle of pereonite 3. Pleopod 2 6, copulatory stylet just reaching to distal mar- 
gin of rami. Uropodal endopod distally broadly rounded, margin with 
rounded teeth; exopod 2.5 times longer than wide, margin with rounded 
teeth, apically obscurely subacute. Posterior margin of pleotelson subtrun- 
cate, with about eight spines. 


RECORDS Jamaica; Puerto Rico; Cozumel, Mexico; Gulf of Mexico. 


Cirolana parva Hansen, 1890 
Figures 59C—E, 60 


DIAGNOSIS 6 6.9mm, 2 7.9 mm. Cephalon with furrow between eyes hav- 
ing middorsal posterior deflection. Antenna reaching posteriorly to pereonite 
4. Pleopod | 6, endopod narrowed in distal third. Pleopod 2 6, copulatory 
stylet reaching by '!/s its length beyond rami. Uropodal endopod with mesial 
margin convex, apically acute; exopod 2.5 times longer than wide, apically 
acute. Pleotelson evenly tapering to angled posterior margin, with seven or 
eight spines. 

RECORDS North and South Carolina; Turks and Caicos Islands; St. 
Thomas and St. Croix, U.S. Virgin Islands; Andros Island, Bahamas; Pu- 


erto Rico; Jamaica; Florida Keys; Dry Tortugas; Barbados; Carrie Bow Cay, 
Belize; Cozumel, Mexico; Panama; Gulf of Mexico; intertidal to 55 m. 


Figure 59. Cirolana obtruncata: A, 2; B, uropod. Cirolana parva: C, 2; D, uropod; E, 
pleotelson and uropods. 


Haptolana 137 


Figure 60. Cirolana parva, 


anteroventral cephalon. 


Creaseriella Rioja, 1953 


DIAGNOSIS Antennular peduncle of two articles (articles 1 and 2 fused). 
Antennal peduncle of five articles. Pereopods all ambulatory. Penial rami 
fused to form short stout process. Pleopod 2 in d with copulatory stylet inser- 
ted at base of endopod. Pleopods 3—5, endopods lacking marginal setae. 
Pleonite 5 lacking free lateral margins. 


Creaseriella anops (Creaser, 1936) 
Figure 61A,B 


DIAGNOSIS 6 15.5 mm. Animal able to conglobate. Eyes lacking. Frontal 
lamina pentagonal, longer than wide, with transverse ridge at widest point. 
Maxillipedal endite with four or five coupling hooks. Pleotelson wider than 
long, posterior margin very broadly rounded and finely crenulate. Both 
uropodal rami distally rounded, margins bearing spines and setae, sympod 
produced along mesial margin of endopod. 


RECORDS Several caves and cenotes on the Yucatan Peninsula, Mexico. 


Haptolana Bowman, 1966 


DIAGNOSIS Mandibular palp directed posteriorly. Antennular peduncle of 
two articles, basal article expanded. Pereopods all prehensile, with dactyli 
closing in propodal groove. Pleopods 3—5, exopods with partial suture and 
marginal setae; endopods undivided, lacking marginal setae. Pleonite 4 over- 
lapping pleonite 5 laterally. 


138 FLABELLIFERA * CIROLANIDAE 


Figure 61. Creaseriella anops: A, adult; B, frontal lamina. Haptolana trichostoma: C, 
adult; D, frontal lamina. Oncilorpheus stebbingi: E, juvenile (from Paul and Menzies, 
1971). 


Haptolana trichostoma Bowman, 1966 
Figure 61C,D 


DIAGNOSIS 6 13.8 mm. Eyes lacking. Frontal lamina broad, pentagonal, 
with anterior angle very broad. Pleotelson wider than long, roughly rec- 
tangular, posterior margin faintly crenulate, with spine between and setae on 
crenulations. Uropodal endopod distally broad, exopod distally narrowly 
rounded, 2/3 width of endopod, both rami with marginal setae and spines. 


RECORDS Cave in Camaguey Province, Cuba. 


REMARKS A second species of Haptolana, H. somala Messana and Chelazzi, 
has been described from northern Somalia in Africa. 


Natatolana 139 


Oncilorpheus Paul and Menzies, 1971 


DIAGNOSIS’ Frontal lamina projecting ventrally. Pleopod 1, exopod indu- 
rate, opercular; endopod membranous, less than half width of exopod. 
Uropodal sympod longer than rami, slightly produced along mesial margin 
of endopod; rami inserted subapically. 


Oncilorpheus stebbingi Paul and Menzies, 1971 
Figure 61E 


DIAGNOSIS @ 11.0 mm. Body narrow, about five times longer than wide. 
Pleotelson triangular, bearing faint middorsal longitudinal ridge, apex nar- 
rowly rounded. 


RECORDS Off Venezuela, 73 m. 


Subfamily Conilerinae, new name 


DIAGNOSIS Clypeus flattened. Frontal lamina flat, narrow. Antennal ped- 
uncular articles 3 and 4 subequal. Pereopods 1-3, ischium and merus pro- 
duced anterodistally. Pereopods lacking secondary unguis on dactyli. Nata- 
tory setae present on pereopods 4—7. 


REMARKS In a discussion of the Cirolanidae, Botosaneanu, Bruce, and 
Notenboom (1986:412) refer to the subfamilies Eurydicinae and Cirolaninae 
but place the Conilera group under the heading “Unnamed Subfamily.” For 
consistency, the Conilera group is here recognized as a subfamily. 


Key to genera of Conilerinae 


1. Uropodal endopod with distal notch; pereopods 4—7 natatory, ischium, 

MEKUS: ANGUCATOUS MAUCENEM) 2.5 cons. cates eek note oe oes Politolana 
Uropodal endopod lacking distal notch; pereopods 4—7 with basis 

flattened and expanded, bearing natatory setae .......... Natatolana 


Natatolana Bruce, 1981 


DIAGNOSIS Frontal lamina narrow; clypeus flat. Pereopods 1—3 bearing 
long setae. Pleopod 2 6, copulatory stylet articulating basally on endopod. 


140 FLABELLIFERA ¢ CIROLANIDAE 


Natatolana gracilis (Hansen, 1890) 
Figure 62 


DIAGNOSIS. 6 8.0 mm. Antennule short, not reaching distal end of antennal 
peduncle. Antenna reaching posteriorly to pereonite 4. Pleopod 2, copulatory 
stylet of endopod cylindrical, bowed, distally rounded. Pleotelson with ob- 
tuse apex, with slight transverse indentation near anterior margin. 


RECORDS Probably St. Thomas, U.S. Virgin Islands; off Sombrero Light, 
Florida, 100—120 m. 
Northern Brazil, 7-85 m. 


REMARKS Hansen (1890) indicated some uncertainty about the exact type 
locality, but thought it likely to have been St. Thomas. Koening (1972) re- 
corded this species from several localities off northern Brazil, but did not 
illustrate her material. 


Politolana Bruce, 1981 


DIAGNOSIS Frontal lamina slender, flattened; clypeus flattened. Antennal 
peduncular articles 3—5 subequal. Pereopods 1—3 with ischium and merus 
anterodistally produced. Pereopods 4—7 with ischium, merus, and carpus 
flattened. 3d pleopod 2 with copulatory stylet inserted basally on endopod. 
Endopod of pleopod 5 lacking marginal setae. Pleonite 5 overlapped laterally 
by pleonite 4. Uropodal endopod with distal emargination; exopod slender, 
elongate; sympod produced along mesial margin of endopod. 


Key to species of Politolana 


1. Uropodal endopod broad distal to emargination, margin obliquely 
truncate; coxae of pereonites 2—6 with impressed line ...... impressa 
Uropodal endopod distal to emargination somewhat narrowed, margin 
evenly convex; coxae of pereonites lacking impressed line ..... polita 


Politolana impressa (Harger, 1883) 
Figure 63A,B 


DIAGNOSIS. 6 and 2 up to 27 mm. Frontal lamina slightly expanded ante- 
riorly. Coxae of pereonites 2—6 with impressed oblique line. Uropodal endo- 


Figure 62. Natatolana gracilis: A, 2; B, frontal lamina; C, pleotelson; D, uropod; E, 
pereopod 7. 


142 FLABELLIFERA * CIROLANIDAE 


\ 


ok 
a 
f 


\ F 


Figure 63. Politolana impressa: A, 3, lateral view; B, uropod, (setae and spines 
omitted). Politolana polita: C, 3; D, pereopod 1; E, uropod, setae and spines 
omitted; F, pereopod 1. 


pod broad distal to emargination, margin subtruncate. Pleotelson posteriorly 


broadly rounded. 


RECORDS’ Massachusetts to Palm Beach, Florida, 32—650 m. 


Politolana polita 143 


Figure 64. Politolana polita, 
anteroventral cephalon. 


Politolana polita (Stimpson, 1853) 
Figures 63C—F, 64 


DIAGNOSIS 6 27.0 mm, 2 29.0 mm. Antennule barely reaching distal end 
of antennal peduncle. Frontal lamina basally slender, anteriorly expanded, 
just visible in dorsal view. Coxae lacking impressed oblique line. Uropodal 
endopod distal to emargination narrowed, margin convex. Pleotelson pos- 
teriorly broadly rounded. 


RECORDS Bay of Fundy, Canada, to Florida Keys, 2—600 m; Gulf of 
Mexico. 


Subfamily Eurydicinae Stebbing, 1905 


DIAGNOSIS Clypeus projecting. Pleonite 5 with free lateral margins (except 
in Xylolana). Penes prominent. Pleopod 2 of d with copulatory stylet artic- 
ulating subbasally, medially, or subapically. 


Key to genera of Eurydicinae 


1. Uropodal sympod not produced along mesial margin of endopod 
Eurydice 
Uropodal sympod produced along mesial margin of endopod ....... 2 


2. Rostrum prominent, fused with frontal lamina, separating antennal 
SES aie Bole hy SES COS eo ae RO tors ree RE EOS ee ree Try ar 5 


(continued ) 


144 FLABELLIFERA * CIROLANIDAE 


Key to genera of Eurydicinae (Continued) 


3: -Pleon: of five ireepleanites. plus pleotelson .).. 2... ...<=- soeae oe wees 4 
Pleon of three free pleonites plus pleotelson ......:...20%:- Colopisthus 


4. Endopods of pleopods 3—5 lacking marginal setae, or with no more 
than three marginal setae; copulatory stylet of endopod of pleopod 2 


in; ©, articulating SUB emmmMANy (oc... sc see see occ we ess Arubolana 
Marginal setae lacking only on endopod of pleopod 5; copulatory stylet 

of pleopod 2 endopod in ¢ articulating basally ........ Metacirolana 

5. Clypeus conical; uropodal endopod lacking notch in outer distal margin 

tee Sal BRE APSR wt aR Ny st ae PREG 7a emt alee cage ce Xylolana 
Clypeus flattened; uropodal endopod with notch in outer distal margin 

gin aE AMPS ees Saran cs kus Wy Syeras a MONG aes eae SURES ara ie Oi il a Excirolana 


Arubolana Botosaneanu and Stock, 1979 


DIAGNOSIS Animal not able to conglobate. Blind, or with very small eyes. 
Anterior margin of frontal lamina broad. Antennular peduncle of three arti- 
cles. Maxillipedal palp of four articles (articles 2 and 3 fused). Maxilla 2 
reduced, endite unarmed, exopod with few marginal setae. Pereopods | and 
2 and sometimes pereopod 3 prehensile; pereopods 4—7 ambulatory. 
Pleopods | and 2, rami undivided. Pleopod 2 3 with copulatory stylet artic- 
ulating subterminally on endopod. Pleopods 3-5, exopods biarticulate; endo- 
pods lacking marginal setae, or with few setae on endopods of 3 and 4; 
pleopod 5 exopod with marginal setae interrupted. 


Key to species of Arubolana 
1. Byes: present, Salk yiycta 33.2 GUN AR ee parvioculata 
VCS. ASETIE a0) car. tayay'e' arcs at a ae a crea a ee caren 2 


2. Pleotelson posteriorly rounded; rostrum not distinct in dorsal view 


BS TAS Chaka te See ay ee ee aruboides 


Arubolana parvioculata 145 


Arubolana aruboides (Bowman and Iliffe, 1983) 
Figure 65A—C 


DIAGNOSIS 6 3.9mm, 2 4.1 mm. Body about three times longer than wide. 
Eyes absent. Antennular peduncle article 3 longer than articles 1 and 2 to- 
gether. Antenna reaching posteriorly to pereonite 6 or 7. Frontal lamina vis- 
ible in dorsal view between antennal bases, anteriorly rounded and only 
slightly wider than proximally, with distally flared ridge on ventral (exposed) 
surface. Pleotelson as long as basal width, evenly narrowing to broadly 
rounded posterior margin, latter with few small serrations and setae. Uropo- 
dal exopod four times longer than wide; endopod length about twice basal 
width, distally obliquely truncate, with elongate setae on inner margin. 


RECORDS Church Cave and Wonderland Cave, Bermuda. 


Arubolana imula Botosaneanu and Stock, 1979 
Figure 65D 


DIAGNOSIS. ¢ and 2 6.25 mm. Body 2.3 times longer than wide. Eyes ab- 
sent. Antenna reaching posteriorly to pereonite 4 or 5. Rostrum distinct, 
anteriorly truncate, separating antennal bases, fused with rectangular frontal 
lamina ventrally. Pleotelson basally slightly wider than long, posterior mar- 
gin broadly rounded to subtruncate, with irregular crenulations or faint 
teeth. Uropodal exopod apically acute, reaching to about midlength of endo- 
pod; latter distally broad, with slight tooth at distolateral angle. 


RECORDS Mangel Cora Tunnel, Aruba. 


Arubolana parvioculata Notenboom, 1984 
Figure 65E,F 


DIAGNOSIS 6 2.8 mm, ¢ 2.9 mm. Body 3.3 times longer than wide. 
Cephalon with tiny pigmented eyes. Antenna reaching posteriorly to per- 
eonite 5. Frontal lamina projecting, dorsally visible. Pleotelson basally wider 
than long, tapering to broadly rounded/subtruncate posterior margin bear- 
ing about six low teeth. Uropodal exopod distally acute, almost three times 
longer than basal width; endopod distally serrate, apically acute. 


RECORDS Interstitial water near Discovery Bay, Jamaica. 


146 FLABELLIFERA * CIROLANIDAE 


Figure 65. Arubolana aruboides: A, 3; B, pereopod 1; C, pereopod 2. Arubolana imula: 
D, 2. Arubolana parvioculata: E, 3; F, pleopod 2, ¢. 


Colopisthus Richardson, 1902 


DIAGNOSIS Cephalon broader than long, becoming triangular between an- 
tennal bases. Pleon consisting of three short free pleonites (often obscured 
beneath pereonite 7), plus triangular pleotelson. 


Eurydice convexa 147 


Colopisthus parvus Richardson, 1902 
Figure 66A 


DIAGNOSIS @ 3.6 mm. Eyes large, well pigmented. Frontal lamina prox- 
imally narrow, anteriorly widened to truncate distal margin. Antennules and 
antennae short, latter reaching posteriorly to pereonite 1. Pleotelson with 
strong middorsal ridge. 


RECORDS’ Bermuda; Puerto Rico, intertidal rocks and algae. 


Eurydice Leach, 1815 


DIAGNOSIS Antennular peduncle article 2 at right angle to article 1. Anten- 
nal peduncle of four articles. Frontal lamina usually slender; clypeus usually 
a ventrally directed triangular blade. Maxillipedal endite reduced, lacking 
coupling hooks. ¢ pleopod 2 with copulatory stylet articulating at mid- 
length. Pleopod 5, endopod lacking marginal setae. Pleonite 5 with free lat- 
eral margins, not overlapped by pleonite 4. Uropodal sympod not produced 
along medial margin of endopod. 


Key to species of Eurydice 


imenrontal lamina distally truncate to faimtly bilobed .........5...-.-. z 
Frontal lamina lanceolate, distally acute: 2.52 e 4-2 2 chan: personata 


2. Posterior margin of pleotelson between notches rounded, with four 
INOdEerAbe SPIES! a. Shree ee HEE EE ake was Deis P98 convexa 
Posterior margin of pleotelson between notches almost straight, with 
four very short spines and several elongate setae ........... piperata 


Eurydice convexa Richardson, 1900 
Figures 66B—E, 67A,B 


DIAGNOSIS. 6 6.1 mm, 2 6.1 mm. Frontal lamina slender, anteriorly widen- 
ing slightly and becoming truncate to faintly bilobed. Posterior margin of 
pleotelson between lateral notches convex, with four spines and few setae 
between serrations; spines between three and four times longer than wide. ¢d: 
Plicate process on antennal flagellar articles about '/s length of article. 
Pleopod 2 with copulatory stylet distally blunt, reaching well beyond rami. 


148 FLABELLIFERA * CIROLANIDAE 


Figure 66. Colopisthus parvus: A, 2. Eurydice convexa: B, 3; C, uropod; D, antennule; 
E, pleotelsonic apex. Eurydice personata: F, 3, lateral view; G, pleotelsonic apex. 
Eurydice piperata: H, pleotelsonic apex. 


RECORDS South Carolina to Florida Keys; Bahamas; Gulf of Mexico and 
Caribbean. 


REMARKS Whether E. convexa and E. littoralis are conspecific needs further 
investigation. Differences can be detected in the male plicate organs of the 
antennae, and in the mandibular palp spination, but range of variation in 


Excirolana 149 


these features is still unknown. It seems unlikely that the species recorded as 
E. littoralis by Moreira (1972) from Brazil is the same species. 


Eurydice personata Kensley, 1987a 
Figures 66F,G; 67C 


DIAGNOSIS <6 6.0 mm, ovigerous 2 5.1—6.4 mm. Frontal lamina slender, 
lanceolate, anteriorly acute. Posterior margin between notches faintly con- 
vex, with four relatively elongate spines (inner pair five or six times longer 
than wide) and few setae between dentitions. ¢: Plicate organ on antennal 
flagellar articles half length of article. Pleopod 2, copulatory stylet on endo- 
pod clavate, barely reaching beyond ramus. 


RECORDS Bermuda; off Georgia, 18—27 m; off South Carolina, 34 m; off 
Miami, Florida; Puerto Rico, 13-17 m; Bahamas, 1—2 m and surface 
plankton tow; Haiti; Cuba; Venezuela. 


REMARKS ‘This recently discovered species has masqueraded under the 
names of E. convexa and E. littoralis for some time, which may explain some of 
the inconsistencies in the literature, especially in variation in the pleotelsonic 
apex. 


Eurydice piperata Menzies and Frankenberg, 1966 
Figure 66H 


DIAGNOSIS 6 4.0 mm, ? 4.5 mm. Frontal lamina slender, widening ante- 
riorly to become slightly bilobed. Posterior margin of pleotelson between 
notches straight to faintly convex, with four spines barely twice longer than 
wide, and several much longer setae between dentition. d: Plicate organ on 
antennal flagellar articles about '/eé length of article but situated subdistally. 
Pleopod 2, copulatory stylet clavate, reaching well beyond ramus. 


RECORDS Georgia to Florida, Gulf of Mexico, 37-150 m. 


Excirolana Richardson, 1912a 


DIAGNOSIS Cephalon with prominent rostrum separating antennular bases; 
fused with flattened frontal lamina. Clypeus with short, broadly triangular 
blade projecting anteroventrally. Antennal peduncle with four or five arti- 
cles. Maxillipedal endite with single coupling hook. Pleopods 3—5, endopods 


150 FLABELLIFERA * CIROLANIDAE 


Figure 67. Eurydice convexa: A, anteroventral cephalon; B, frontal lamina. Eurydice 


personata: C, frontal lamina. 


lacking marginal setae. Pleonite 5 with free lateral margins, not overlapped 
by pleonite 4. Uropodal sympod produced along mesial margin of endopod. 


Key to species of Excirolana 


1. Pleotelson with two anterior hollows clearly joined by impressed line; 


uropodal endopod about half length of exopod .......... braziliensis 
Pleotelson with two anterior hollows not connected by impressed line; 
uropodal endopod about two-thirds length of exopod ....... mayana 


Excirolana braziliensis Richardson, 1912a 
Figures 68A—C, 69A—C 


DIAGNOSIS 6 6.0 mm, 2 7.5 mm. Frontal lamina very slender between 
antennal bases, widening anteriorly into rounded structure between anten- 


VY 


.?) 


Figure 68. Excirolana braziliensis: A, 6; B, pleopod 2 6; C, uropod, (setae and 
spines omitted). Excirolana mayana: D, 2; E, pleopod 2 ¢; F, uropod, setae and 
spines omitted. 


152 FLABELLIFERA * CIROLANIDAE 


Figure 69. Excirolana braziliensis: A, anteroventral cephalon; B, pleotelson; C, 


uropodal endopod. Excirolana mayana: D, anterodorsal cephalon; FE, anteroventral 
cephalon. 


nular bases, joined to rostrum by very slim isthmus. Clypeus distally sub- 
acute. Uropodal endopod about half length of exopod. Pleotelson with two 
lateral hollows defined and connected by clear impressed line; posterior mar- 
gin evenly convex, bearing numerous plumose setae. 


Metacirolana {533 


RECORDS Caribbean to Brazil; common in the intertidal of sandy beaches; 
Gulf of Mexico. 
Gulf of California to Chile. 


REMARKS Glynn et al. (1975) produced a thorough study of the taxonomy, 
zonation, and distribution of this Pan-American species. 


Excirolana mayana (Ives, 1891) 
Figures 68D-—F, 69D,E 


DIAGNOSIS 6 8.2mm, 2 10.0 mm. Frontal lamina between antennal bases 
about half anterior width. Clypeus anteriorly rounded. Uropodal endopod 
about 7/3 length of exopod. Pleotelson with two faint lateral hollows in ante- 
rior half, not connected by impressed line. 


RECORDS Florida to Venezuela, intertidal. 


Metacirolana Nierstrasz, 1931 


DIAGNOSIS. Frontal lamina anteriorly dilated, free, projecting; clypeus tri- 
angular, projecting ventrally. Maxillipedal endite with one coupling hook. 
Pleon with five free segments, pleonite 5 not overlapped laterally by pleonite 
4. Eyes often larger, and antennular flagellum of more articles in d than in 


a 


Key to species of Metacirolana 


cts OME POSECHOE Ly LEUMEALC oho ances 4 >, s ckepoyeye ePeiahs josie dis: Afo Shale xs halia 
Melson posteriorly rounded Or angulate . ... . o..<cjs.sjoi ois ieee eis. > wiecarar 2 
2. Posterfr margin of telson an obtuse angle ................ agaricicola 
SCEScrta OS LELIOU yah OUNCE sn ote a0e ais: bos vine sie eto bi © (ob lap eh es nse ae rete 3 


3. Posterior margin of telson narrowly rounded; uropodal rami, margins 
SEM OITGUI A CUCIMEAEG eri Pela, Stan ee Oe aia nl ace lb yes cvelays waist menziest 
Posterior margin of telson broadly rounded; uropodal rami, margins 
BBSOULELY Cente x. seis On eee Lie. Sr hace sphaeromiformis 


154 FLABELLIFERA * CIROLANIDAE 


Metacirolana agaricicola Kensley, 1984 
Figure 70A—C 


DIAGNOSIS d 2,6 mm, ovigerous 2 2.1 mm. Antennular flagellum of six or 
seven articles. Antennal flagellum of 10 articles. Posterior margin of telson 
finely dentate, with broadly obtuse median point. Uropodal exopod about 
half width of endopod, margins dentate, apically acute; endopod, margins 
dentate, distally angled, apically acute. 


RECORDS Carrie Bow Cay, Belize, 1-20 m, in coral on reef slope, and spur 


and groove zone. 


Metacirolana halia Kensley, 1984 
Figure 70D-F 


DIAGNOSIS 6 2.9 mm, ovigerous 2 2.7 mm. Antennular flagellum of 10 
articles in 2, 14 in d. Antennal flagellum of 10 articles in 2, 11 in ¢. Pos- 
terior margin of telson truncate, bearing about eight sensory spines. Uropo- 
dal exopod distally broadly rounded, more than half distal width of endopod, 
outer margin dentate, with about 11] sensory spines; endopod distally broad, 
margin straight, bearing about 12 sensory spines. 


RECORDS Carrie Bow Cay, Glover’s Reef, Belize; intertidal to 23 m; Turks 
and Caicos Islands, 1 m; Bahamas; Jamaica; Cozumel, Mexico. 


Metacirolana menziesi Kensley, 1984 
Figure 71A,B 


DIAGNOSIS 6 2.3 mm, ovigerous 2 2.4 mm. Antennular flagellum of six 
articles in 9, eight in 6. Antennal flagellum of nine articles in 2, 10 in 6. 
Posterior margin of telson broadly rounded, finely dentate. Uropodal exopod 
half width of endopod, margins dentate, apically acute; endopod, margins 


dentate, apically acute. 


RECORDS Carrie Bow Cay, Belize, intertidal to 30 m, usually in coral 
rubble. 


Metacirolana sphaeromiformis (Hansen, 1890) 
Figure 71C,D 


DIAGNOSIS. 6 2.5 mm, 2 3.2 mm. Antennular flagellum of three articles. 
Antennal flagellum of eight articles. Posterior margin of telson narrowly 


Metacirolana sphaeromiformis 193 


Figure 70. Metacirolana agaricicola: A, 2; B, uropod; C, maxilliped. Metacirolana 
halia: D, uropod; E, 3; F, &. 


rounded, obscurely dentate. Telson with low rounded middorsal ridge and 
pair of lateral ridges. Uropodal exopod more than half width of endopod, 
margin dentate; exopod distally broadened, margin dentate, with few sen- 
sory spines. 


156 FLABELLIFERA * CIROLANIDAE 


Figure 71. Metacirolana menziesi: A, 2; B, uropod. Metacirolana sphaeromiformis: C, 2; 
D, uropod. 


RECORDS Looe Key, Florida, intertidal reef crest; Turks and Caicos Is- 
lands, | m; St. Thomas, U.S. Virgin Islands. 


Xylolana Kensley, 1987a 


DIAGNOSIS Frontal lamina and rostrum fused, broad, separating antennu- 
lar bases. Clypeus conical, projecting. Maxillipedal endite reduced, lacking 


Alcirona iay7/ 


coupling hooks; palp of five articles. Copulatory stylet in d articulating in 
distal half of mesial margin of pleopod 2 endopod. Pleopods 3—5, exopods 
biarticulate; endopods lacking marginal setae. Pleonite 5 lacking free lateral 
margins, overlapped by pleonite 4. Uropodal sympod produced along mesial 
margin of endopod. 


Xylolana radicicola Kensley, 1987a 
Figure 72 


DIAGNOSIS 6 2.6mm, 2 3.3 mm. Body about four times longer than great- 
est width. Uropodal exopod about */3 width of endopod, bearing single short 
subapical spine; both uropodal rami distally rounded. Pleotelson with lateral 
margins subparallel, with poorly defined middorsal longitudinal ridge. 


RECORDS ‘Twin Cays, Belize, in dead red mangrove roots, | m. 


Family Corallanidae Hansen, 1890 


DIAGNOSIS Outer ramus of maxilla 1 apex with single strong falcate spine, 
with single strong spine with one or more smaller hooked spines at base, or 
with two large recurved spines, occasionally with one to three smaller spines 
between them. Maxillipedal endite reduced or lacking. 


Key to genera of Corallanidae 


ee Maxilla | with single strong falcate spime. 2.20. 2/60) .13% es ee ee sa soe 2 
Maralla 1, with: two large talcate:spines: «. 24. :2) Pejsn. w ser ivi 2 Alcirona 


2. Maxilla 2, distal article slender-elongate; article 2 of maxillipedal palp 


COVES S| ae rae thelial suse Seer Nite tp ia aera ete 1 pew Nalicora 
Maxilla 2 distally bluntly bilobed; article 3 of maxillipedal palp longest 
soc, GRUPPE gees ae ae ay, Pate ais arena rote ater Ota Excorallana 


Alcirona Hansen, 1890 


DIAGNOSIS Antennular peduncle of two articles. Mandible lacking molar. 
Maxilla | having two large recurved spines, with one or more smaller spines 
between these. Maxilla 2 a simple rounded lobe. Maxilliped lacking endite. 
Posterior half of body hirsute. 


158 FLABELLIFERA * CORALLANIDAE 


Key to species of Alcirona 


1. Golden-brown setae starting dorsally on pereonite 3; pereopod 1, 
dactylus having several elongate spines ................... insularis 
Golden-brown setae starting dorsally on pereonites 5 or 6; pereopod 1, 
dactylus having accessory spine jonly @. 2... 10.0.4 46. c eee krebsi 


Alcirona insularis Hansen, 1890 
Figure 73A 


DIAGNOSIS. 6 and @, 5.0 mm. Posterior half of body, especially in d, bear- 
ing stiff golden-brown setae, these beginning as posterior row on pereonite 3, 
and becoming dense on posterior pereonites, pleonites and pleotelson. Per- 
eopods 1-3, dactylus strongly falcate, having distal unguis equal in length to 
rest of article, and with three or four strong teeth on posterior margin. Apex 
of pleotelson rounded, bearing six short marginal spines in addition to setae. 


RECORDS Looe Key, Florida, 0.5—-6 m; St. Thomas, U.S. Virgin Islands, 
40—46 m; Puerto Rico, from intertidal coral rubble and from gills of nurse 
shark Ginglymostoma sirratum; St. Lucia, from coral rubble. 


Alcirona krebsii Hansen, 1890 
Figure 73B—D 


DIAGNOSIS <6 10 mm, ovigerous 2 15.5 mm. Posterior half of body, espe- 
cially ind bearing stiff golden-brown setae, these beginning in posterior row 
on pereonites 5 or 6, becoming dense on posterior pereonites, pleonites, 
uropods, and pleotelson. Pereopod 1, dactylus strongly falcate, with unguis 
equal in length to rest of article, and with one strong tooth and several low 
tubercles on posterior margin. Apex of pleotelson rounded, bearing six short 
spines in addition to setae. 


RECORDS’ Bermuda, in sponges; Florida Keys; Quintana Roo, Yucatan 
Peninsula; St. Thomas, U.S. Virgin Islands; Venezuela. 


REMARKS A single 8-mm ¢ specimen of Alcirona from Panama Bay (Pa- 
cific) has the characteristic pereopod | of A. krebsii, but has the rows of stuff 
setae beginning on about pereonite 3. The possibility that A. krebsii is another 
amphi-Panamic species needs to be investigated. 


Excorallana 159 


Figure 72. Xylolana radicicola: A, B, 3; C, maxilliped; D, pleopod 2 d; E, pleopod 3. 


Excorallana Stebbing, 1904 


DIAGNOSIS Eyes well developed and pigmented, sometimes contiguous or 
nearly so. Maxilla 1, outer ramus a single falcate spine. Maxillipedal palp of 
five articles; endite reduced or absent. Pereopods 1—3 subprehensile or pre- 


160 FLABELLIFERA « CORALLANIDAE 


Figure 73. Alcirona insularis: A, pereopod 1. Alcirona krebsi: B, 3; C, maxilla 1; D, 
pereopod 1. 


hensile, pereopods 4—7 ambulatory. All rami of pleopods bearing marginal 
setae. Pleotelson often with characteristic spination and tuberculation; lat- 
eral margins often with incision. 


REMARKS Excorallana subtilis (Hansen, 1890) was described from St. 
Thomas, U.S. Virgin Islands, based on a specimen undergoing ecdysis; the 
true identity of this species remains uncertain. 

In the key, two species have been included which are not illustrated. These 
are E. mexicana Richardson, 1905, from the Gulf of Mexico, and E. delaneyi 


Excorallana 161 


Stone and Heard, 1989, from the northeastern Gulf of Mexico. The latter 
species, particularly, could conceivably be encountered in the Florida Keys. 

Delaney (1984) provides a useful review of the genus Excorallana, and of the 
distribution of the species. 


Key to species of Excorallana 


MPP ESAGOULIPUOUSE. |. Haste eck clas qslstettiihk « Jul+ siertintis ® Gate ae 5 Deas 2 
Eves Well SGpabdateted.. shisha, os at Aeids, 4 Unie. eden Me eM eee 4 
Beepex of plcotclson with deep slit .. ..\...Jcnusisglees co oe eee fissicauda 
Pese MaMa PUCOLEISONSCMENS. y.2 02 cos Stacie tanta s ae «i ceeimioe S Oe EO oe es 3 
pepelecotcison-with lateral ancision:.......05.. i. 52 2. eas she Gees » oculata 
Picoteisonmacking lateral incision: ~ . .< = a)..s is. see sas 6 eee warmingit 
Peercotcison with lateral incision” =. o 6.0. 3. ko tees neces es 6 
Picoteison lacking fateral incision’. <2. soso eee ea wee yess es 5 
5. Frontal lamina linguiform, anteriorly rounded ............. berbicensis 
Frontal lamina posteriorly with faintly concave margins, anteriorly 
SELEY KS he Metre Ce Oe PRE PUR ere MEP mee APENAYIBN Meng GEA delaneyi 
6. Frontal lamina strongly grooved for entire length ................. 7 
Frontal lamina with,ventral surface flat. )s< i.cca.... fs.0 22 250s ncn 8 


7. Pleotelson with medial row of small tubercles flanked by row of larger 
Pubercles may postenon Wale 7.362) 2) no o1 od cepa slaeiaw es mexicana 
Pleotelson lacking rows of tubercles in posterior half ........ antillensis 


8. Frontal lamina distinctly bell shaped; d cephalon with two pairs of 
tubercles and antennular bases not tuberculate ......... quadricornis 
Frontal lamina, and 6 cephalon and antennules not as above ....... 9 


9. Frontal lamina anteriorly broadly rounded, length 1.5 or less times 
basal width; d cephalon with two pairs of tubercles and basal 
antennulararticle each with tubercle: <.c.. .1. <:cr.6 os: 9 sisiam are sexticornis 

Frontal lamina anteriorly narrowly rounded, length about twice basal 
width; ¢d cephalon with three tubercles, tubercles lacking on 
AACIEINULALMOASES! “(Sec URE sa: Sais etatoe nls oleae sere s tricornis tricornis 


162 FLABELLIFERA * CORALLANIDAE 


Excorallana antillensis (Hansen, 1890) = E aculicauda (Miers 18 I) 
Figure 74A—D 


DIAGNOSIS. 6 11.0 mm, ovigerous 2 15.0 mm. Cephalon unornamented. 
Eyes well separated. Frontal lamina parallel sided, length twice basal width, 
anteriorly broadly rounded. Pleonites 2—4, posterolateral margins tubercu- 
late, middorsal posterior margin excavate with strong middorsal tubercle; 
pleonite 5 with strongest tubercles submedian, posterior margin not exca- 
vate. Pleotelson with low middorsal ridge, strong basal tubercles; lateral inci- 
sions present; two submedian patches of spines; apex narrowly rounded. 


RECORDS Florida Keys; St. Thomas, U.S. Virgin Islands; Puerto Rico; 
Quintana Roo, Mexico; Carrie Bow Cay, Belize, 5—18 m. 


Excorallana berbicensis Boone, 1918 
Figure 74E,F 


DIAGNOSIS 6 9.9 mm, 2 12.0 mm. Eyes well separated. Frontal lamina 
about twice longer than wide, posteriorly parallel sided, widening anteriorly 
to broadly rounded apex. Subadult 6 cephalon unornamented except for 
pair of very low tubercles mesial to eyes. Posterior margins of pleonites 3—5 
very faintly tuberculate. Pleotelson with submedian pair of low tubercles 
basally; lacking lateral incisions; apex rounded. 


RECORDS Guyana; French Guiana; Guadeloupe. 


Excorallana fissicauda (Hansen, 1890) 


DIAGNOSIS 11 mm. Cephalon unornamented. Eyes contiguous. Frontal 
lamina unknown. Pleonite 5 with three strong mesial, and several smaller 
lateral tubercles on posterior margin. Pleotelson with two strong submedian 
basal tubercles; lateral incision lacking; posterior margin with deep open 
incision. 

RECORDS St. Thomas, U.S. Virgin Islands. 


REMARKS This species was described from a single specimen, and has not 
been recorded since. After examining the holotype, Paul Delaney (in litt.) 
suspects that the terminal incision of the pleotelson may be the result of an 
injury. 


Excorallana oculata 163 


F 


Figure 74. Excorallana antillensis: A, 6; B, frontal lamina; C, maxilla 2; D, 
maxilliped. Excorallana berbicensis: E, 3; F, frontal lamina. 


Excorallana oculata (Hansen, 1890) 
Figure 75A,B 


DIAGNOSIS 6 6.9 mm, 2 8.5 mm. Eyes contiguous. Cephalon unorna- 
mented. Frontal lamina slender, linguiform, widest posteriorly. Pleonites 3—5 
each with slightly hollowed middorsal area containing strong flattened tuber- 
cle; pleonite 5 with two strong flanking tubercles; pleotelson basally with low 
median ridge and two strong submedian tubercles; short strong spines in two 
roughly triangular submedian patches; lateral incisions present; apex nar- 
rowly rounded. 


RECORDS’ Bahamas; Cuba, Puerto Rico, 40 m; Barbados. 
Brazil. 


Figure 75. Excorallana oculata: A, 3; B, frontal lamina. Excorallana quadricornis: C, 
3; D, frontal lamina. Excorallana sexticornis: E, 36; F, frontal lamina. Excorallana 
tricornis tricornis: G, 3; H, frontal lamina. Excorallana warmingi: I, 3; J, frontal 


lamina. 


Excorallana tricornis tricornis 165 


Excorallana quadricornis (Hansen, 1890) 
Figures 75C,D; 76A—C 


DIAGNOSIS 6 13.2 mm, 2 12.1 mm. Eyes well separated. Cephalon in 6 
with two pairs of tubercles, anterior pair connected by low rounded ridge, 
posterior pair situated mesial to eyes. Frontal lamina bell shaped, broadest 
posteriorly. Pereonite 1 with submedian pair of low tubercles. Posterior mar- 
gin of pereonite 7 and pleonites faintly tuberculate. Pleotelson with two sub- 
median raised areas bearing short spines; lateral margins with incision; few 
low tubercles basally. 


RECORDS Bermuda; St. Thomas, U.S. Virgin Islands; Jamaica, intertidal 
in grassflats and between mangrove roots; Martinique; Belize; Venezuela. 


Excorallana sexticornis (Richardson, 1901) 
Figures 75E,F; 76D-—F 


DIAGNOSIS 6 7.9mm, ovigerous 2 6.9—8.3 mm. Eyes well separated. Basal 
antennular peduncular article in d with short anterodorsally directed tuber- 
cle. Cephalon with two pairs of prominent tubercles, anterior pair shorter 
than posterior pair, latter situated mesial to eyes. Frontal lamina, length less 
than twice width, sides parallel to faintly converging anteriorly, apically 
broadly rounded. Posterior margins of pleonites 2—5 with low rounded tuber- 
cles, those near middorsal line largest. Pleotelson with two basal submedian 
tubercles, numerous scattered dorsal spines, lateral margins with incision, 
apex narrowly rounded. 


RECORDS Key West, Florida; Cuba; Puerto Rico; Twin Cays, Belize, shal- 
low infratidal from dead mangrove wood. 


Excorallana tricornis tricornis (Hansen, 1890) 
Figures 75G,H; 77 


DIAGNOSIS 6 8.2 mm, 2 9.9 mm. Basal antennular article narrow, not 
dilated. Cephalon in d with one median and two dorsal “horns.” Eyes large, 
well separated. Frontal lamina between two and three times longer than 
basal width, sides subparallel, anteriorly rounded to subacute. Pereon 
smooth. Margins of lateral incision of pleotelson separated by gap; anterior 
margin of incision lined with short spines; scattered short spines on dorsum 
of telson, but especially concentrated in two submedian patches. Uropodal 
exopod, length 2.3—2.5 times width; apical notch nearly symmetrical. 


166 FLABELLIFERA * CORALLANIDAE 


400HM 


Figure 76. Excorallana quadricornis: A, cephalon and pereonite 1 6; B, ventral 
cephalon; C, frontal lamina enlarged. Excorallana sexticornis: D, cephalon and 
pereonite 1 6; E, cephalon @; F, ventral cephalon. 


RECORDS’ Turks and Caicos Islands, 1 m; St. Thomas and St. Croix, U.S. 
Virgin Islands, 48—55 m; Cuba; Puerto Rico, on gills of rays Aetobatus narinari 
and Dasyatis americana, and on squirrel fish; Belize, intertidal to 15.2 m, in 
intertidal coral rubble, in coarse sediments in Syringodium and Thalassia sea- 
grass beds, on brown alga Turbinaria, on Madracis sp. sponge, on Agaricia sp. 
coral; Gulf of Mexico. 


Excorallana warmingii 167 


Figure 77. Excorallana tricornis tricornis: A, cephalon and pereonite 1 6; B, ventral 


cephalon; C, pleotelson and uropods. 


REMARKS The subspecies Excorallana tricornis occidentalis Richardson, 1905a, 
from southern California, differs from the Gulf and Caribbean subspecies in 
lacking a gap between the margins of the pleotelsonic incision, and in having 
a relatively wider uropodal exopod which shows a distinctly asymmetrical 
apical notch. 


Excorallana warmingii (Hansen, 1890) 
Figure 751, J 


DIAGNOSIS 6 9.7 mm, ¢ 12.0 mm. Cephalon unornamented. Eyes con- 
tiguous, occupying most of dorsal surface of head. Posterior margins of 
pleonites very faintly tuberculate. Frontal lamina, length slightly more than 
twice basal width, tapering anteriorly to rounded apex. Pleotelson unorna- 
mented except for two faint submedian tubercles basally; lateral incisions 
lacking; apex broadly rounded, with five low but distinct marginal teeth. 


168 FLABELLIFERA * CORALLANIDAE 


Ra 


— 


“— 


Figure 78. Nalicora rapax: A, 2; B, maxilla 1; C, maxilla 2; D, maxilliped. 


RECORDS Bahamas; between Cuba and the Yucatan Pensinsula; Puerto 
Rico. 
Off Brazil near Rio de Janeiro. 


Nalicora Moore, 1901 


DIAGNOSIS Maxilla 1 exopod a single strongly falcate distal spine with 
knoblike mesial process, and basal caplike convex papilla-bearing structure. 
Maxilla 2 of four articles, distal article slender. Maxillipedal palp of five 
articles; endite lacking. 


FLABELLIFERA * CYMOTHOIDAE 169 


Nalicora rapax Moore, 1901 
Figure 78 


DIAGNOSIS 6 6.9 mm, ovigerous 2 10.0 mm. Eyes well developed. Frontal 
lamina basally slender, widening anteriorly, apex subacute. Posterior half of 
body bearing numerous scattered stiff setae. Pereonites 4—7 with row of low 
rounded tubercles near posterior margin. Posterior margins of pleonites 3-5 
faintly tuberculate, more noticeable in d. Pleotelson wider than long; lateral 
margins faintly sinuous; apex rounded. 


RECORDS Florida Keys, 55 m; Puerto Rico, 50-150 m; Gulf of Mexico off 
lorida, 37—73 m. 


Family Cymothoidae Leach, 1818 


DIAGNOSIS. Antennules and antennae reduced, no clear distinction between 
peduncles and flagella. Mandibular palp of three articles. Maxilla 1 with four 
terminal spines. Maxilla 2 apically bilobed, armed with several spines. Max- 
illipedal palp of two articles, terminal article bearing hooks. All seven pairs of 
pereopods prehensile, ending in strongly hooked dactyli. Pleopods lacking 
marginal setae in adults. 


REMARKS The cymothoids are exclusively ectoparasites on marine, fresh- 
water, and brackish-water fishes. Most cymothoids occur in shallow water, 
mainly in tropical and subtropical areas. The position of attachment on the 
host (externally, in the buccal cavity, or in the gill chamber) is usually genus- 
or species-specific. The body of gill parasites is often asymmetrical, being 
slightly twisted, perhaps an effect of the position on the host. The mouthparts 
are highly adapted for the parasitic mode of life, while all seven pairs of 
pereopods are strongly prehensile. The posterior pereopods of some genera 
have the basal article expanded and carinate, allowing for increased mus- 
culature. The secretion of anticoagulants in the juvenile stages further aids 
the blood-feeding habit. The surface area of the pleopods is often increased 
by the development of lobes on the bases or the lamellae, providing an in- 
creased respiratory ability. 

The post-mancal juvenile stages (sometimes referred to as the aegathoid 
stage) have large eyes, and highly setose pleopods for active swimming. The 
juveniles will attach themselves indiscriminantly to any convenient fish host, 
but eventually attach to the preferred host-species. The juvenile then de- 
velops into a functional male, losing the swimming setae of the pleopods. 
Both juveniles and males feed actively, drawing blood from the host fish. The 


170 FLABELLIFERA * CYMOTHOIDAE 


Key to genera of Cymothoidae 


. Antennule broader and usually longer than antenna; cephalon very 


weakly sunkiimto. Perconmite, Lo. 05 erect aut x «8-0 the tao ae Ste eee, eee 2 
Antennule not broader or longer than antenna; cephalon distinctly 

immersed in, or not at all immersed in pereonite 1 .............. 4 

2. Bases,of-antennules widely Separated: ioc os ane... oo eee 3 

Bases of antennules comtieuwous: 2.6.4.0. san 4s. e. 7 see: Glossobius 

. Body curved to one side; pleonite 1 extended laterally more on one side 

thanon others ic. a.doa shee ee attic eee ee woe ote Mothocya 
Body rarely curved to one side; pleonite 1 extended equally on each 

SIGE £0 loo can cs pour ne ens Gaetan kam Ae SR Nc mee heen ae Renocila 


ee Le an CON ee Net eee eT ee eee Fae Agarna 


. Cephalon not immersed in pereonite 1; posterior margin of cephalon 
{LISITMATE «0-2 Need nares: 2 ele etn Siete otochae ate eee ee ea aie ee eee 6 
Cephalon to some degree immersed in pereonite 1; posterior margin of 
cephalon ‘not trsinMater 4 tary He = poland daca wie peel uel wise ed 7 


Posterolateral angles of pereonites 2—6 not produced; coxal plates short, 
rarely reaching posterior margin of their pereonites ........ Anilocra 

Posterolateral angles of pereonites 2—6 posteriorly increasingly 
produced; coxal plates usually reaching to posterior margin of their 


PCREORULES. ofa are. cud acters aera) Oa at re EE Ose ea Nerocila 

. Basal antennular articles expanded and contiguous ......... Ceratothoa 
Basal antennular articles expanded but not contiguous, or basal 

antennular articles neither expanded nor contiguous ............. 8 

Basal antennular articles expanded but not contiguous ......... Kuna 

Basal antennular articles neither expanded nor contiguous ......... a 


9. Pleonal margins continuous with pereonal margins, pleon not abruptly 
narrowed, only weakly immersed in pereonite 7 ........... Lironeca 

Pleon to some degree narrower than pereon; pleon usually deeply 
IMUINERSeC AN. PET CONME Jj cos iia eee haat ee ecie ee Cymothoa 


FLABELLIFERA * CYMOTHOIDAE 171 


male eventually becomes a female (all cymothoids are protandrous) should a 


female not already be present. In some species, the female is nonfeeding. In 


those species which settle either in the mouth cavity or gill chamber of the 


host, integumental pigment is frequently lost, and the eyes become reduced. 


Given the highly variable morphology of the cymothoids, in part imposed 


by the parasitic mode of life, and the existence of polymorphism and possible 


sibling species, the taxonomy of this family demands the examination of large 


numbers of specimens. As a further aid to identification, Table 3 is provided, 


giving host species, parasite, and site of attachment. 


TABLE 3. CYMOTHOID PARASITES FROM THE CARIBBEAN AREA, LISTED BY FISH 


HOST SPECIES 


Fish host 


Cymothoid parasite 


Site of attachment 


Abudefduf saxatilis 


Acanthurus bahianus 


Acanthurus chirurgus 


Alutera schoepfi 
Anchoa lamprotaenia 
Apogon lachneri 
Apogon maculatus 
Apogon townsendi 
Arius felis 
Astrapogon stellatus 
Batrachoides surinamensis 
Caranx hippos 
Caranx latus 

Caranx ruber 

Caranx sp. 
Chaetodipterus faber 
Chaetodon capistratus 
Chaetodon ocellatus 
Chaetodon sedentarius 
Chaetodon striatus 


Anilocra abudefdufi 
Kuna insularis 
Anilocra acanthuri 


Anilocra acanthuri 


Nerocila acuminata 
Lironeca tenuistylis 
Mothocya bohlkeorum 
Renocila colini 
Renocila colini 
Nerocila acuminata 
Mothocya bohlkeorum 
Nerocila acuminata 
Cymothoa oestrum 
Cymothoa oestrum 
Cymothoa oestrum 
Cymothoa oestrum 
Nerocila acuminata 
Anilocra chaetodontis 
Anilocra chaetodontis 
Anilocra chaetodontis 
Anilocra chaetodontis 


beneath eye 

gill chamber 

? at base of pectoral fin; 
immature on or near 
pectoral or pelvic fin 

? at base of pectoral fin; 
immature on or near 
pectoral or pelvic fin 

on or at base of fin 

posterior to pectoral fin 

in gill chamber 

next to dorsal fin 

next to dorsal fin 

on or at base of fin 

in gill chamber 

on or at base of fin 

inside mouth 

inside mouth 

inside mouth 

inside mouth 

on or at base of fin 

beneath eye 

beneath eye 

beneath eye 

beneath eye 


(continued ) 


172 


TABLE 3. (Continued) 


Fish host 


Cymothoid parasite 


Site of attachment 


Chilomycterus schoepfi 
Chromis cyaneus 
Chromis multilineatus 
Cynoscion nebulosus 
Cynoscion sp. 
Epinephelus cruentatus 
Epinephelus fulvus 
Epinephelus guttatus 
Epinephelus itajara 
Epinephelus sp. 
Exocoetus spp. 

Gerres rhombeus 
Haemulon aurolineatum 
Haemulon carbonarium 
Haemulon chrysargyreum 
Haemulon flavolineatum 
Haemulon macrostomum 
Haemutlon plumieri 
Haemulon sciurus 


Hemirhamphus brasiliensis 


Hirundichthys speculifer 
Holacanthus tricolor 
Holocentrus ascensionis 


Hyporhamphus unifasciatus 


Leiostomus xanthurus 


Lepiosteus spatula 
Lutjanus analis 
Lutjanus mahogoni 
Lutjanus synagris 
Megalops atlanticus 
Monacanthus ciliatus 
Mugil cephalus 
Mpyripristis jacobus 


Ocyurus chrysurus 
Orthopristis chrysoptera 


Nerocila acuminata 
Anilocra chromis 
Anilocra chromis 
Cymothoa excisa 
Cymothoa oestrum 
Anilocra haemuli 
Anilocra haemuli 
Anilocra haemuli 
Nerocila acuminata 
Cymothoa oestrum 
Glossobius impressus 
Lironeca redmanni 
Anilocra haemuli 
Anilocra haemuli 
Anilocra haemuli 
Anilocra haemuli 
Anilocra haemuli 
Anilocra haemuli 
Anilocra haemuli 
Glossobius hemirhamphi 
Glossobius impressus 
Anilocra holacanthi 
Anilocra holocentri 


Mothocya nana 
Nerocila acuminata 
Cymothoa excisa 
Lironeca redmanni 
Nerocila acuminata 
Cymothoa excisa 
Cymothoa excisa 
Cymothoa excisa 
Cymothoa oestrum 
Nerocila acuminata 
Nerocila acuminata 
Anilocra myripristi 


Cymothoa excisa 
Cymothoa excisa 


on or at base of fin 
beneath eye 
beneath eye 
inside mouth 
inside mouth 
beneath eye 
beneath eye 
beneath eye 
on or at base of fin 
on or at base of fin 
inside mouth 
in gill chamber 
beneath eye 
beneath eye 
beneath eye 
beneath eye 
beneath eye 
beneath eye 
beneath eye 
inside mouth 
inside mouth 
beneath eye 
2 between eyes, d and 
immature beneath 
eye 
in gill chamber 
on or at base of fin 
inside mouth 
in gill chamber 
on at base of fin 
inside mouth 
inside mouth 
inside mouth 
inside mouth 
on or at base of fin 
on or at base of fin 
2 between eyes, imma- 
ture beneath eye 
inside mouth 
inside mouth 


Agarna cumulus 


173 


Fish host 


Cymothoid parasite 


Orthopristis ruber 


Anilocra haemuli 


Site of attachment 


beneath eye 


Paranthias furcifer Anilocra haemuli beneath eye 
Phaeoptyx conklini Mothocya bohlkeorum in gill chamber 
Phaeoptyx pigmentaria Mothocya bohlkeorum in gill chamber 


Pogonias cromis 
Pomacentrus partitus 
Priacanthus arenatus 
Scomberomorus cavalla 
Scomberomorus maculatus 
Scomberomorus regalis 
Selar crumenophthalmus 
Serranus tigrinus 


Sphoeroides maculatus 
Synodus foetens 


Nerocila acuminata 
Anilocra partiti 
Cymothoa oestrum 
Lironeca redmanni 
Lironeca redmanni 
Lironeca redmanni 
Cymothoa oestrum 
Renocila bowmani 
Renocila waldneri 
Nerocila acuminata 
Cymothoa excisa 


on or at base of fin 


beneath eye 
inside nouth 

in gill chamber 
in gill chamber 
in gill chamber 
inside mouth 
next to dorsal fin 
next to dorsal fin 


on or at base of fin 


inside mouth 


Agarna Schioedte and Meinert, 1883 


DIAGNOSIS Cephalon with posterior margin not trilobed; immersed in per- 
eonite 1. Antennular bases contiguous. Pereonites 4—7 on one side flattened 
and expanded; coxal plates of pereopods 4—7 also expanded and flattened 
but generally hidden by lateral expansion of pereonites. Bases of posterior 
three pereopods with well-formed carinae. Pleonites 1 and 2 immersed in 
pereonite 7; pleonites 2—5 with free fingerlike lateral margins. 


Agarna cumulus (Haller, 1880) 
Figure 79 


DIAGNOSIS ¢ 18 mm. Eyes present, indistinct. Pereon strongly “humped” 
dorsally. Uropod about '/3 length of pleotelson; uropodal exopod slightly 
longer, and twice width of endopod. Pleotelson triangular, length */s basal 
width, apex rounded. 


RECORDS No host recorded: Key West, Florida. 


174 FLABELLIFERA * CYMOTHOIDAE 


Figure 79. Agarna cumulus: A, 2, dorsal view; B, °, ventral view, coxal plates 
stippled. 


Anilocra Leach, 1818 


DIAGNOSIS Cephalon usually narrowed anteriorly to triangular apex folded 
ventrally between bases of antennules; posterior margin trilobed; not im- 
mersed, or only weakly immersed in pereonite 1. Coxal plates small, com- 
pact, not reaching level of posterior margin of their respective pereonites. 
Pereopods increasing in length posteriorly, pereopod 7 often markedly longer 
than 6. Pleon not immersed or only slightly immersed in pereonite 7. 
Pleopods 3—5 often formed into deep pockets or pleats. Uropods often ex- 
tending beyond pleotelsonic apex. 


REMARKS Williams and Williams (1981) have provided a comprehensive 
treatment of this genus and nine of its species in the West Indies. Table | in 
this latter paper provides characters for separating these nine species. This 
table also indicates that for each species, the site of attachment of the adult to 
the host fish is specific, with six species attaching under the eye of the host. 


Anilocra abudefdufi 175 


Key to species of Anilocra 


1. Pereopods 2—4 with swelling on outer margin of dactylus ........... 2 
Pereopods 2—4 lacking swelling on outer margin of dactylus ........ 5 
BeBody axis distorted by more than 10° .. 2222508 ea ae. holacanthi 
Paolveasisdistarned by lessthano vii sao: os eee 3 
Be actylus of percopod 7 longer than propodus .:.......'......+.. partiti 
Mactylus of percopod.7 shorter than propodus~).0.5 2.5... <-- 4 
4. Posteroventral angle of pereonite 7 overlapping pleonite | only 
a a ee Ce cyte Pr oe raid catia ede agora ae (epee abudefdufi 
Posteroventral angle of pereonite 7 overlapping pleonites 1 and 2 
sna eer Sk ees teat a7o ese, uausiths ale a Giant chaetodontis 
a rosteroventral angle of pereonite 7 produced ........:). 0.0 c00.505+ 6 
Eestcroventral angle of pereonite 7 not produced .............:.5... 7 
6. Uropod reaching posterior margin of pleotelson ............ myripristis 
Uropod not reaching posterior margin of pleotelson .......... haemuli 
7. Posteroventral angle of pereonite 7 overlapping pleonite 1 .... holocentri 
Posteroventral angle of pereonite 7 not overlapping pleonite 1 ....... 8 
8. Uropod reaching posterior margin of pleotelson ............. acanthun 
Uropod not reaching posterior margin of pleotelson ........... chromis 


Anilocra abudefdufi Williams and Williams, 1981 
Figure 80A—C 


DIAGNOSIS Ovigerous 2 19.0-31.0 mm, ¢ 7.0—8.5 mm. Pereopods 2—4 
with swelling on outer margin of dactylus. Posteroventral angle of pereonite 6 
slightly produced, of pereonite 7 more produced, overlapping pleonite 1. 
Uropodal endopod variable, not reaching, to extending well beyond, apex of 
exopod. Color: upper lateral half to three-fourths of dorsal surface of 2 when 
attached to host is dark brown; rest of dorsal surface light brown to yellow. 
Attaching beneath eye of host. 


RECORDS Sergeant major Abudefduf saxatilis: Panama; Colombia. 


176 FLABELLIFERA * CYMOTHOIDAE 


Figure 80. Anilocra abudefdufi: A, 2, lateral view; B, pereopod 3; C, pereopod 7. 
Anilocra acanthuri: D, pleotelson and uropods. Anilocra chaetodontis: E, 9, lateral 
view. Anilocra chromis: F, pleotelson and uropods. 


Anilocra acanthuri Williams and Williams, 1981 
Figure 80D 


DIAGNOSIS Ovigerous ¢ 29.0—40.0 mm, 6 4.0-8.0 mm. Pereopods 2-4 
without swelling on outer margin of dactylus. Posteroventral angles of per- 


eonites not produced. Uropod not reaching posterior margin of pleotelson. 


Endopod of uropod variable, not reaching, to extending well beyond, apex of 
exopod. Color: dorsal surface of 2 black to lead gray, ventral surface gray. 


Attaching under pectoral fin of host. 


RECORDS Doctorfish Acanthurus chirurgus: Florida Keys; Bahamas; Puerto 
Rico; U.S. Virgin Islands. Ocean surgeon Acanthurus bahianus: Florida Keys; 
Bahamas; Cuba; Jamaica; Dominican Republic; Puerto Rico; U.S. Virgin 


Islands. 


Anilocra haemuli 177 


Anilocra chaetodontis Williams and Williams, 1981 
Figure 80E 


DIAGNOSIS Ovigerous 2? 18-28 mm, d 4—5 mm. Pereopods 2—4 with swell- 
ing on outer margin of dactylus. Posteroventral angles of pereonites 4—7 be- 
coming progressively produced, that of pereonite 7 overlapping pleonite 2. 
Uropod not reaching posterior margin of pleotelson; uropodal endopod ex- 
tending beyond apex of exopod. Pleotelson as wide as long to slightly wider 
than long. Color: dorsal surface of ? black to lead gray, ventral surface gray. 
Attaching beneath eye of host. 


RECORDS Foureye butterflyfish Chaetodon capistratus: Bahamas; Puerto Rico; 
British and U.S. Virgin Islands. Banded butterflyfish Chaetodon striatus: 
Bahamas; Puerto Rico; British Virgin Islands. Spotfin butterflyfish Chaetodon 
ocellatus: Bahamas; Puerto Rico; U.S. Virgin Islands. Reef butterflyfish 
Chaetodon sedentarius: Puerto Rico. 


Anilocra chromis Williams and Williams, 1981 
Figure 80F 


DIAGNOSIS Ovigerous 2 16-28 mm, d 4—9 mm. Pereopods 2-4 lacking 
swelling on outer margin of dactylus. Posteroventral angles of pereonites not 
produced. Uropod extending beyond posterior margin of pleotelson; uropo- 
dal endopod not reaching beyond exopod. Color: upper lateral one-fourth to 
two-thirds of dorsal surface of 2 when attached is dark gray, shading to off- 
white lower lateral area. Attaching beneath eye of host. 


RECORDS Brown chromis Chromis multilineatus: Puerto Rico; British and 
U.S. Virgin Islands. Blue chromis Chromis cyaneus: Bahamas; Dominican Re- 
public. No host recorded: Anguilla. 


Anilocra haemuli Williams and Williams, 1981 
Figure 81A,B 


DIAGNOSIS Ovigerous 2? 21-40 mm, d 7 mm. Body axis distorted less than 
5°. Pereopods 2—4 lacking swelling on outer margin of dactylus. Postero- 
ventral angle of pereonites 6 and 7 produced, latter overlapping pleonite 1. 
Uropod not reaching posterior margin of pleotelson; uropodal endopod 
reaching beyond apex of exopod. Color: dorsal surface of ¢ yellow to light 
brown. Attaching beneath eye of host. 


178 FLABELLIFERA * CYMOTHOIDAE 


Figure 81. Anilocra haemuli: A, 2, dorsal view; B, ¢, lateral view. Anilocra 
holacanthi: C, 2. Anilocra holocentri: D, 2°. Anilocra myripristis: E, pleotelson and 
uropods. Anilocra partiti: F, 2; G, pereopod 7. 


RECORDS French grunt Haemulon flavolineatum: Florida Keys; Puerto Rico; 
British and U.S. Virgin Islands. Tomtate Haemulon aurolineatum: Jamaica; 
Puerto Rico. Smallmouth grunt Haemulon chrysargyreum: Puerto Rico; U.S. 
Virgin Islands. Caesar grunt Haemulon carbonarium: Puerto Rico; U.S. Virgin 


Anilocra holocentri 179 


Islands. Spanish grunt Haemulon macrostomum: Puerto Rico. White grunt 
Haemulon plumieri: Florida Keys; Yucatan Peninsula. Bluestriped grunt 
Haemulon sciurus: Florida Keys. Cora cora Orthopristis ruber: Margarita Island, 
Venezuela. Coney Epinephelus fulvus: Bahamas; Dominican Republic; Puerto 
Rico; U.S. Virgin Islands; Guadeloupe. Red hind Epinephelus guttatus: Puerto 
Rico; British and U.S. Virgin Islands. Graysby Epinephelus cruentatus: 
Bahamas; Dominican Republic; U.S. Virgin Islands. Creole-fish Paranthias 
furcifer: Dominican Republic; Puerto Rico; Colombia. No host recorded: 
Cuba; Jamaica; Dominica; Barbados; Venezuela; Brazil. 


Anilocra holacanthi Williams and Williams, 1981 
Figure 81C 


DIAGNOSIS Ovigerous 2 21-33 mm, 6 4—7 mm. Body axis distorted by 
more than 10°. Pereopods 2—4 with swelling on outer margin of dactylus. 
Posteroventral angles of pereonites 5—7 progressively more produced, that of 
pereonite 7 overlapping pleonite 1. Uropod not reaching posterior margin of 
pleotelson; uropodal endopod reaching beyond apex of exopod. Color: dorsal 
surface of 2 black to lead gray. Attaching beneath eye of host. 


RECORDS Rock beauty Holacanthus tricolor: Bahamas; Jamaica; Dominican 
Republic; Puerto Rico; British and U.S. Virgin Islands. 


Anilocra holocentri Williams and Williams, 1981 
Figure 81D 


DIAGNOSIS Ovigerous 2 32—46 mm, ¢ 5-9 mm. Body axis distorted less 
than 5°. Pereopods 2—4 lacking swelling on outer margin of dactylus. 
Posteroventral angle of pereonite 7 produced, overlapping pleonite 1. 
Uropod not reaching posterior margin of pleotelson; uropodal endopod 
reaching beyond apex of exopod. Color: dorsal surface of 2 dark brown, 
ventral surface light brown. @ attaching between eyes of host; d or transi- 
tional stage beneath eye. 


RECORDS Squirrelfish Holocentrus ascensionis: Puerto Rico; U.S. Virgin 
Islands. 
No host recorded: Patagonia, Straits of Magellan. 


180 FLABELLIFERA * CYMOTHOIDAE 


Anilocra myripristis Williams and Williams, 1981 
Figure 81E 


DIAGNOSIS Ovigerous 2 29-40 mm, ¢ 6—7 mm. Body axis distorted less 
than 5°. Pereopods 2—4 lacking swellings on outer margin of dactylus. 
Posteroventral angle of pereonites 6 and 7 produced, latter overlapping 
pleonite 1. Uropod reaching beyond posterior margin of pleotelson; uropodal 
endopod reaching beyond apex of exopod. Color: dorsal surface of 2 light 
reddish brown, ventral surface yellow. 2 attaching between eyes of host; 
immature or transitional forms sometimes beneath eye. 


RECORDS Blackbar soldierfish Myripristis jacobus: Bahamas; Dominican Re- 
public; Puerto Rico. 


Anilocra partiti Williams and Williams, 1981 
Figure 81F,G 


DIAGNOSIS Ovigerous 2 12—16 mm, transitional 7.6—9.0 mm. Body axis 
distorted less than 5°. Pereopods 2—4 with swelling on outer margin of dac- 
tylus. Pereopod 7 with dactylus longer than propodus. Posteroventral angle 
of pereonite 7 produced, overlapping pleonite 1. Uropod not reaching pos- 
terior margin of pleotelson; uropodal endopod not reaching apex of exopod. 
Color: dorsal surface black to slate gray. Attaching beneath eye of host. 


RECORDS Bicolor damselfish Pomacentrus partitus: Jamaica. 


Ceratothoa Dana, 1852 


DIAGNOSIS Cephalon more or less immersed in pereonite 1, posterior mar- 
gin not trisinuate. Bases of antennules expanded, contiguous. Coxal plates 
compact; anterior plates not extending beyond posterior margins of their 
respective pereonites; posterior coxal plates may or may not be produced 
beyond the posterior margins of the pereonites. Anterior pleonites narrowed, 
immersed in pereonite 7. Copulatory stylet lacking on pleopod 2 of d of some 
species. 


Ceratothoa deplanata Bovallius, 1885 
Figure 82A 


DIAGNOSIS 2 18 mm. Cephalon subtriangular, anterior margin rounded. 
Pereopods 4—7 with strongly carinate bases. Uropod reaching or extending 


Ceratothoa deplanata 181 


Figure 82. A, Ceratothoa deplanata (from Bovallius, 1885); B, Cymothoa caraibica; C, 
Cymothoa excisa; D, Cymothoa oestrum. 


slightly beyond posterior margin of pleotelson; rami subequal in length and 
width. Pleotelson basally wider than long, posterior margin broadly 
rounded. Color: bright yellow. 


RECORDS Haiti, host not recorded. 


182 FLABELLIFERA * CYMOTHOIDAE 


Cymothoa Fabricius, 1793 


DIAGNOSIS Body usually not distorted. Cephalon with posterior margin not 
trilobed; more or less immersed in pereonite 1; latter with anterolateral cor- 
ners produced to embrace cephalon. Bases of antennules not expanded, well 
separated. Anterior coxal plates not reaching posterior borders of their re- 
spective pereonites, posterior coxal plates nearly reaching or extending be- 
yond posterior borders of pereonites. Pleon narrower than, and immersed in 
pereonite 7. Pleonites increasing in length and width posteriorly. 


Key to species of Cymothoa 


1. Anterolateral angles of pereonite 1 reaching to half length of cephalon 


on lesseyesior tracest@l eyes present (ass 25. 52a oe, oem ete 2 
Anterolateral angles of pereonite 1 broad, reaching to anterior margin 

of cephalons-eyesvalsemt: ii. /e ca ate path cena ie ou sere Sofas oestrum 

2. Anterolateral angles of pereonite 1 narrow, subacute ........... excisa 

Anterolateral angles of pereonite 1 broad, rounded .......... caraibica 


Cymothoa caraibica Bovallius, 1885 
Figure 82B 


DIAGNOSIS ¢ 17 mm, d 12—16 mm. Anterior margin of cephalon broadly 
rounded. Eyes large, distinct. Broadly rounded anterolateral angles of per- 
eonite 1 reaching to about midlength of cephalon. Bases of pereopods 4—7 
with strong, rounded carina. Uropodal rami subequal in length, equal to 
peduncle in length. Pleotelson width about twice length, posterolateral mar- 
gin broadly rounded. 


RECORDS Puerto Rico; Gulf of Mexico. 


Cymothoa excisa Perty, 1833 
Figure 82C 


DIAGNOSIS Ovigerous 2 20-24 mm. Anterior margin of cephalon in dorsal 
view truncate to slightly excavate; eyes small, indistinct. Anterolateral angles 
of pereonite 1 narrowly rounded to subacute, reaching anteriorly to about 
midlength of cephalon. Pereopods 4—7 with high rounded carina on basis. 


Glossobius 183 


Uropods hardly reaching halfway along lateral margin of pleotelson; exopod 
slightly longer than endopod. Pleotelson about twice wider than long; 
broadly rounded and somewhat bilobed. 


RECORDS Yellowtail snapper Ocyurus chrysurus: Yucatan Peninsula, Mexico; 
Carrie Bow Cay, Belize; Margarita Island, Venezuela; Panama. Mutton 
snapper Lutjanus analis: Yucatan Peninsula, Mexico; Panama. Lane snapper 
Lutjanus synagris: Panama. Mahogany snapper Lutjanus mahogoni: Panama. 
Pigfish Orthopristis chrysoptera: Florida, Gulf of Mexico. Spot Leiostomus 
xanthurus: Texas, Gulf of Mexico. Spotted seatrout Cynoscion nebulosus: Texas, 
Gulf of Mexico. Inshore lizardfish Synodus foetens: Texas, Gulf of Mexico. No 
host recorded: Massachusetts; South Carolina; Georgia; Florida Keys; 
Bahamas; Cuba; Trinidad; Brazil. 


Cymothoa oestrum (Linnaeus, 1793) 
Figure 82D 


DIAGNOSIS Ovigerous 2 38 mm. Cephalon in dorsal view with ante- 
rolateral angles rounded, anterior margin slightly excavate; eyes absent. An- 
terolateral angles of pereonite 1 expanded, broadly rounded, reaching to 
level of anterior margin of cephalon. Pereonites 4—7 with high rounded 
carina on basis. Uropod reaching posteriorly beyond midlength of 
pleotelson; exopod slightly longer than endopod. Pleotelson length slightly 
more than half basal width. 


RECORDS Bigeye scad Selar crumenophthalmus: Bermuda; U.S. Virgin Is- 
lands. Bigeye Priacanthus arenatus: Bermuda. Bar jack Caranx ruber: Florida 
Keys; Carrie Bow Cay, Belize. Horse-eye jack Caranx latus: Bahamas; Bar- 
bados. Crevalle jack Caranx hippos: Venezuela. Jack Caranx sp.: Jamaica; Cu- 
racao. Hind Epinephelus sp.: Grenada. Parrotfish: Jamaica. Seatrout Cynoscion 
sp.: Panama. Tarpon Megalops atlantica: Texas, Gulf of Mexico. No host re- 
corded: Honduras; Haiti. 


Glossobius Schioedte and Meinert, 1883 


DIAGNOSIS Cephalon not immersed in pereonite 1; excavate on either side 
in anterior half, forming broad and anteriorly rounded median area; anten- 
nae fitting into excavate areas. Bases of antennules contiguous, expanded. 
Antennules broader and longer than antennae. Bases of pereopods 4—7 with 
posterior margin expanded and flattened. Pleonites 1-3 immersed in per- 
eonite 7. 


184 FLABELLIFERA * CYMOTHOIDAE 


Key to species of Glossobius 


1. Coxal plates of pereonites 1 and 2 anteroventrally protruding impressus 
Coxal plates of pereonites 1 and 2 close to body, not protruding 
a Nin a en tee aac apa er cae NOME So cat sr eer a, A aes hemirhamphi 


Glossobius hemiramphi Williams and Williams, 1985a 
Figure 83A 


DIAGNOSIS Ovigerous ¢ 27 mm. Eyes small but distinct. Cephalon pointed 
anteriorly. Fused coxa of pereonite | and free coxa of pereonite 2 carinate but 
not protruding. Coxa of pereonite 7 semicircular in dorsal view. Pleotelson 
with middorsal length more than half basal width; lateral margins somewhat 
tapered; posterior margin variable, sinuate or excavate. Uropods reaching to 
or slightly beyond posterior pleotelsonic margin; rami subequal in length, 
exopod slightly broader than endopod. 


RECORDS Ballyhoo Hemiramphus brasiliensis: Puerto Rico. 


Glossobius impressus (Say, 1818) 
Figure 83B 


DIAGNOSIS Ovigerous ¢ 33 mm. Eyes small but distinct. Cephalon 
rounded anteriorly. Fused coxal plate of pereonite | and distinct coxal plate 
of pereonite 2 protruding strongly in oblique anteroventral direction. Uropod 
reaching to posterior half of pleotelson; exopod shorter and narrower than 
endopod. Pleotelson basal width twice length, posteriorly broadly bilobed. 
Attaching inside mouth of host. 


RECORDS Flyingfish Exocoetus spp.: Rio de Janeiro, Brazil; North Atlantic, 
especially in the Gulf Stream. 

Mirrorwing flyingfish Hirundichthys speculifer: North Atlantic. No host rec- 
ord: Senegal, West Africa. 


Kuna Williams and Williams, 1986 


DIAGNOSIS Cephalon somewhat immersed in pereonite 1. Anterior margin 
of pereonite | not trisinuate. Number of articles in antennules and antennae 


Figure 83. A, Glossobius hemiramphi; B, Glossobius impressus; C, Kuna insularis; D, 
Lironeca redmani; E, Lironeca tenuistylis. 


reduced. Antennule somewhat expanded; basal article expanded but not 
contiguous. Copulatory stylet present on pleopods 1—3 in d. Pleonites dor- 
sally strongly convex, not immersed in pereonite 7. 


186 FLABELLIFERA * CYMOTHOIDAE 


Kuna insularis (Williams and Williams, 1985b) 
Figure 83C 


DIAGNOSIS Ovigerous 2? 11.1-17.2 mm, 6 4.2—8.7 mm, transitional 9.6— 
9.8 mm. Antennules and antennae consisting of four articles each. Uropods 
short, not reaching posterior margin of pleotelson. Clavate copulatory stylet 
present on pleopods 1-3 in 6. Pleotelson basally broader than long, pos- 
terior margin broadly rounded. 


RECORDS’ Sergeant major Abudefduf saxatilis: Carrie Bow Cay, Belize; Cura- 
cao; Panama. 


Lironeca Leach, 1818 


DIAGNOSIS Cephalon weakly to deeply immersed in pereonite 1; posterior 
border rarely trisinuate. Bases of antennules not expanded, well separated. 
Posterior pereopods with carinae on bases in Gd, carinae present or absent in 
2. Pleonites subequal in width; pleonites 1 and 2 rarely narrowed and 
weakly to moderately immersed in pereonite 7. Pleopods highly folded, and 
with lamellar or digitiform accessory gills in some species. 


Key to species of Lironeca 


1. Uropodal endopod about twice longer than wide; pleon somewhat 


iinInIerSed Nin pErE OM: 2s eee << eee eles sibs ameter gel eae e redmannt 
Uropodal endopod about three times longer than wide; pleon barely 
iIMmMersed ul. PerEOn wt 5.4 oie aes one eee ee tenuistylis 


Lironeca redmanni Leach, 1818 
Figure 83D 


DIAGNOSIS Ovigerous ¢ 19.5—25.0 mm. Cephalon barely immersed in per- 
eonite 1. Pleon somewhat immersed in pereon, but lateral margins of pleonite 
1 free. Pleotelson basally wider than long. Uropodal rami reaching well be- 
yond posterior margin of pleotelson; exopod longer than endopod, both rami 
somewhat broad, endopod about twice longer than wide. Attaching to gills of 
host. 


Mothocya 187 


RECORDS New Jersey to Florida; gills of kingfish, Jamaica; Cuba; St. 
Christopher; Spanish mackerel Scomberomorus maculatus and cero Scomberomorus 
regalis, Puerto Rico; king mackerel Scomberomorus cavalla, Colombia; Gerres rho- 
mbeus, Panama; spot Leiostomus xanthurus, Gulf of Mexico. 

Brazil. 


Lironeca tenuistylis (Richardson, 1912b) 
Figure 83E 


DIAGNOSIS 2 13 mm. Cephalon barely immersed in pereonite 1. Uropodal 
rami reaching beyond rounded posterior margin of pleotelson; exopod longer 
than endopod; endopod slender, about three times longer than wide. Pleonite 
1 barely immersed in pereonite 7. Pleotelson basally wider than long. Attach- 
ing to host between pectoral and anal fin. 


RECORDS Longnose anchovy Anchoa lamprotaenia: Panama. 


Mothocya Costa, 1851 


DIAGNOSIS Cephalon more or less immersed in pereonite 1. Bases of anten- 
nules widely separated; antennules longer and more robust than antennae. 
Coxae nearly reaching or extending beyond posterior margin of respective 
pereonites. Pleon somewhat immersed in pereonite 7. Uropodal exopod 
longer than endopod. 


REMARKS Bruce (1986b) revised the genus Mothocya. The species of Moth- 
ocya are almost entirely gill parasites on the fish families Hemiramphidae, 
Apogonidae, Belonidae, and Atherinidae. 


Key to species of Mothocya 


1. Cephalon anteriorly narrowed, slightly immersed in pereonite 1; 
plentelsonisulsrectamelan) 22 2. fe c.2' oe deo age os wns an bohlkeorum 
Cephalon anteriorly broad, deeply immersed in pereonite 1; pleotelson 
SIOE RIAN UIA ese cre he ee TGCS 2 ci Sige g hs ayers | waar @ see win nana 


188 FLABELLIFERA * CYMOTHOIDAE 


Mothocya bohlkeorum Williams and Williams, 1982 
Figure 84B 


DIAGNOSIS Ovigerous 2 7.6—8.5 mm, d¢ 3.7 mm. Cephalon anteriorly nar- 
rowed in dorsal view, ventrally flexed, broadly rounded; slightly immersed in 
pereonite 1. Pleotelson subrectangular. Uropods extending slightly beyond 
posterior margin of pleotelson; exopod only slightly longer than endopod. ¢ 
lateral lobes of pleopodal peduncles not developed. Endopods of pleopods 3— 
5 with small proximomedial lobe. 


RECORDS Whitestar cardinalfish Apogon lachneri: Puerto Rico. Dusky car- 
dinalfish Phaeoptyx pigmentaria: Bahamas. Freckled cardinalfish Phaeoptyx con- 
klini: Florida Keys; Bahamas. Conchfish Astrapogon stellatus: Leeward 
Islands. 


Mothocya nana (Schioedte and Meinert, 1884) 
Figure 84A 


DIAGNOSIS Ovigerous 2 11.0-17.0 mm, d 7.9—8.3 mm. Cephalon deeply 
immersed in pereonite 1; rostrum anteroventrally narrowly rounded. Uropo- 
dal exopod markedly longer than endopod. Pleotelson broad, with posterior 
margin rounded sufficiently to give appearance of being subtriangular. 


RECORDS Halfbeak Hyporhamphus unifasciatus: Chesapeake Bay, Maryland; 
Georgia; Florida; Colon, Panama. Halfbeak Hemiramphus bermudensis: 
Bermuda. 


Nerocila Leach, 1818 


DIAGNOSIS Body generally more depressed than in most cymothoid genera, 
rarely curved. Cephalon with anterior margin convex, narrowly rounded, or 
concave; not, or only slightly, immersed in pereonite 1. Pereonite 1 anterior 
margin trisinuate. Posterolateral angles of pereonites weakly to strongly pro- 
duced, increasing in length posteriorly. Coxal plates prominent, usually al- 
most reaching or extending to posterior margin of their respective pereonites. 
Juveniles and 6 usually with spines on posterior pereopods; ? lacking these 
spines. Pleon not immersed in pereonite 7. Pleonites subequal in length; 
pleonites 1 and 2 usually produced posterolaterally. Pleopods typically with 
small lamellar accessory gills; pleopods 3—5 folded into deep pockets or 
pleats. Uropods usually extending beyond pleotelsonic apex. 


190 FLABELLIFERA * CYMOTHOIDAE 


Nerocila acuminata Schioedte and Meinert, 1881 


DIAGNOSIS Ovigerous 2 16.2—19.0 mm. Cephalon with anterior margin 
convex. Posterolateral angles of all, or of posterior pereonites only, produced 
into acute or subacute angles. 


RECORDS Striped burrfish Chilomycterus schoepfi: Texas, Gulf of Mexico. 
Northern puffer Sphoeroides maculatus: New York. Striped mullet Mugil 
cephalus: Texas, Gulf of Mexico. Jewfish Epinephelus itajara: Texas, Gulf of 
Mexico. Hogfish: Bermuda. Alligator gar Lepisosteus spatula: Louisiana, Gulf 
of Mexico. Hardhead catfish Arius felis: Texas, Gulf of Mexico. Sawfish: Flor- 
ida (Atlantic). Black drum Pogonias cromis: Texas, Gulf of Mexico. Orange 
filefish Alutera schoepfi: Texas, Gulf of Mexico. Toadfish Batrachoides sur- 
inamensis: Colon, Panama. Spot Leiostomus xanthurus: Florida, Gulf of Mexico. 
Spadefish Chaetodipterus faber: Florida, Gulf of Mexico; Virginia. Fringed fil- 
efish Monacanthus ciliatus: Florida, Gulf of Mexico. No host recorded: Mas- 
sachusetts; Florida Keys; Florida, Gulf of Mexico. Louisiana, Gulf of Mex- 
ico. Texas, Gulf of Mexico. 


REMARKS Brusca (1981) has shown that this highly variable species occurs 
on both sides of the Isthmus of Panama, in two relatively distinct forms. 
Intergrades between the two forms do occur but are uncommon. Brusca 
(1981:159) also lists all the host-records for this species in the eastern Pacific. 


Nerocila acuminata Schioedte and Meinert, 1881, forma acuminata 
Figure 84C 


DIAGNOSIS Cephalon width equal to or greater than length; frontal margin 
narrowly rounded. Posterolateral angles of anterior pereonites weakly pro- 
duced, rounded to subacute; of posterior pereonites more strongly produced, 
subacute to acute. Coxal plates 3—7, 4—7, or 5—7 with acute posterolateral 
angles; coxae rarely reaching beyond posterior margins of their respective 
pereonites. 


Nerocila acuminata Schioedte and Meinert, 1881, forma aster 
Figure 84D 


DIAGNOSIS Cephalon always wider than long; anterior margin broadly 
rounded. Posterolateral angles of all pereonites strongly produced, acute, all 
reaching well beyond posterior margins of their respective pereonites. Coxal 
plates 2—7 strongly produced with acute posterior angles. 


Renocila colini 191 


Renocila Miers, 1880 


DIAGNOSIS’. Body rarely curved. Cephalon anteriorly weakly to distinctly 
truncate. Antennular bases well separated. Antennules and antennae some- 
what flattened, antennules usually broader and longer than antennae. Per- 
eonites 5—7 with posterolateral corners more or less strongly produced. 
Pleonites not laterally incised. 


REMARKS Williams and Williams (1980) provide a key to nine species of 


Renocila. 
Key to species of Renocila 
1. Posteroventral angle of pereonite 7 reaching pleonite 1 .......... colini 
Posteroventral angle of pereonite 7 reaching beyond pleonite 1 ...... Zs 


2. Dorsal surface of body brown; posteroventral angle of pereonite 7 


heaemine pleamite 6 Et chse a oc ed iaje a hint seopeete eta su0 wi/acas waldneri 
Dorsal surface of body black; posteroventral angle of pereonite 7 
REACHING PICOMILEY Oph 2.56 or epee so Se es el Stn tonsa Bg enn edt a mies bowmani 


Renocila bowmani Williams and Williams, 1980 
Figure 85A 


DIAGNOSIS 2 18.0 mm, d 11.5 mm. Posteroventral angles of pereonites 5— 
7 produced, that of pereonite 7 overlapping pleonites 1—3. Pereopods 1—3 
lacking swelling on dactylus. Pereopods 6—7 subequal in length. Uropodal 
exopod longer than endopod. Pleotelson length 3/1 basal width. Color: dorsal 
surface of body and appendages uniform black. Attached to dorsum of body 
close to dorsal fin. 


RECORDS Harlequin bass Serranus tigrinus: Dominican Republic. 


Renocila colini Williams and Williams, 1980 
Figure 85B,C 


DIAGNOSIS Ovigerous 2 12.0—-17.5 mm, ¢6 7.5—-13.0 mm. Pereonites 5—7 
with posteroventral angle produced, that of pereonite 7 overlapping pleonite 
1 only. Pereopods 1—3 lacking swelling on dactyli; pereopods 6—7 subequal in 
length. Uropod reaching beyond pleotelson, endopod more than half length 


192 FLABELLIFERA * CYMOTHOIDAE 


Figure 85. A, Renocila bowmani. Renocila colini: B, 2; C, 3. D, Renocila waldneri. 


of exopod. Pleotelson '/7 to '/2 wider than long, with slight rounded apex. 
Color: dorsal surface of body and appendages uniformly yellowish brown. 
Attached to dorsum of body, close to dorsal fin. 


FLABELLIFERA ¢ LIMNORIIDAE 193 


RECORDS Flamefish Apogon maculatus: Puerto Rico. Belted cardinalfish Ap- 
ogon townsendi: Puerto Rico. 


Renocila waldneri Williams and Williams, 1980 
Figure 85D 


DIAGNOSIS Ovigerous 2 15.3-19.3 mm, d 5.0—-10.8 mm. Posteroventral 
angle of pereonite 5 moderately produced, of pereonites 6—7 more strongly 
produced, that of pereonite 7 overlapping pleonites 1 and 2. Pereopods 1—3 
without swelling on dactyli. Pereopods 6 and 7 subequal in length. Uropodal 
exopod slightly longer than endopod. Pleotelson basally wider than long; 
posterior margin broadly and evenly rounded. Color: dorsal surface of body 
uniform brown; appendages yellowish brown. Attached to dorsum of body 
close to dorsal fin. 


RECORDS Harlequin bass Serranus tigrinus: Dominican Republic. 


Family Limnoriidae Harger, 1879 


DIAGNOSIS Body ovate in cross section, often becoming more setose posteri- 
orly. Cephalon subspherical, freely articulating with pereonite 1; eyes lateral. 
Antennules and antennae well separated at bases. Mandible with strong in- 
cisor; lacking molar and well-defined lacinia mobilis, but with species- 
distinctive lacinioid bristle or seta; palp usually of three articles. Maxillipe- 
dal palp of five articles; endite well developed. Coxae present on pereonites 
2—7. Pleon consisting of five free pleonites plus pleotelson; latter subcircular, 
set obliquely to axis of body, usually with anterolateral crests. Uropod with 
strong protopod inserted ventrolaterally. 


Key to genera of Limnoriidae 


PUropodal rami very unequal 7. 02022 22 22 tees Seek se oe aes 2 
Mropodal rami-subedqtal fet 5 Sas Sh OS oa Paralimnoria 
2. Mandibular incisors possessing rasp and file ................ Limnoria 


Mandibular incisors lacking rasp and file ............... Phycolimnoria 


194 FLABELLIFERA ¢ LIMNORIIDAE 


REMARKS This family includes a number of species that are of considerable 
economic importance. Given that species of Limnoria are wood borers, 
wooden structures such as wharf pilings that are immersed in sea water and 
even in water of reduced salinity are vulnerable to attack by these gribbles. 
Prolonged exposure can lead to weakening and eventual collapse of these 
structures (see Ray, 1959). Even creosote-treated wood is not fully protected; 
Limnoria tuberculata will bore into such wood to where the creosote has not 
penetrated. 

The isopods rasp at the wood fibres with the rasp and file structures of the 
mandibles, usually following the grain of the wood. With this boring activity, 
saprophytic fungi and bacteria invade the wood and assist in the breakdown 
process. Limnoria lack cellulase-secreting microflora in their gut, but proba- 
bly secrete a cellulase themselves (Boyle and Mitchell, 1978). It is also prob- 
able that the fungi and bacteria, the latter often densely aggregated on the 
setae of the isopod, form part of the animals’ diet. In the natural environ- 
ment, Limnoria perform an important role in the breakdown of dead wood, 
especially in mangrove areas. 

Sexual dimorphism of the pleotelson does occur in some species. This as- 
pect of the morphology, however, has hardly been investigated. 


Limnoria Leach, 1814 


DIAGNOSIS Antennular flagellum of four articles. Antennal flagellum of 
three to five articles. Incisor of right mandible equipped with filelike struc- 
ture on upper surface; incisor of left mandible with rasplike structure. Rami 
of pleopod 5 lacking marginal setae. Uropodal exopod much shorter than 
endopod, bearing terminal claw. Pleotelson smooth, or variously ornamented 
with tubercles and ridges. 


Limnoria indica Becker and Kampf, 1958 
Figure 86A,B 


DIAGNOSIS 6 3.0mm, ovigerous 2 3.0 mm. Pleonite 5 with submedian pair 
of strong rounded ridges, converging slightly posteriorly. Pleotelson basally 
with two pairs of submedian tubercles and pair of lateral tubercles. 


RECORDS Cozumel, Mexico; Man o’War Cay, Belize. 
India; Hong Kong; Philippines; east coast of Australia. 


Limnoria insulae 195 


Key to species of Limnoria 


1. Dorsal surface of pleotelson lacking prominent tubercles, ridges, or 
carinae (L. simulata may appear to lack ornamentation; in this 
Species thie tubercles|are very smiall)ri owes ob 1s cbs ee onto om 2 2 

Dorsal surface of pleotelson bearing tubercles, ridges, or carinae .... 3 


2. Pleotelson flat; pleonite 5 with broadly rounded middorsal ridge 


4 cpg Gatch a ie ROP Cue Ro RE RS a a platycauda 
Pleotelson cup shaped; pleonite 5 with strong narrowly rounded 

TINIE EE O/RSE1| Fi) oF Sc oa care are ae aR insulae 

3. Pleotelson with basal tubercles but lacking ridges ................. 4 

Pleotelson with ridges but lacking freestanding tubercles ........... 7 

4. ¢ pleotelson with single strong middorsal tubercle .......... unicornis 

Pleotelson with more than one basal tubercle .......4...¢..22s%... 5 

5 Pleoteisom with: three basal tubercles: ~ 4). s2 66s Obes sok tuberculata 

Pleotelson with more than three basal tubercles ................... 6 


6. Pleotelson with four basal tubercles in line (difficult to detect) simulata 


aupleoteison with six, basal’ tubercles 2208 aieety 5 ee ee ake indica 
7. Pleotelson with single middorsal longitudinal ridge ...... multipunctata 
Plecotelsonswithytwo rounded: basal ridges) 7.22 ye:-/..0 62. syed eis Seale 8 
oe Pleonite) with strong. V-shaped ridge 23... scajed 6 c.c ae os 2 hens pfefferi 
Pleonite 5 with two posteriorly converging ridges .......... saseboensis 


Limnoria insulae Menzies, 1957 
Figure 86C 


DIAGNOSIS’ d6 3.0 mm, ovigerous ¢ 3.4 mm. Pleonite 5 with strong middor- 
sal ridge. Pleotelson cup shaped, lateral crests extended anteromesially, sep- 
arated basally by distinct gap; posterior margin and lateral crests not 
tuberculate. 


RECORDS ‘Twin Cays, Belize. 
Fiji; Guam; Palmyra Island; Caroline Islands. 


196 FLABELLIFERA * LIMNORIIDAE 


Figure 86. Limnoria indica: A, pleotelson, 3; B, pleotelson, 2. Limnoria insulae: C, 
pleotelson. Limnoria multipunctata: D, pleotelson in oblique-lateral view. 


Limnoria multipunctata Menzies, 1957 
Figures 86D; 87A 


DIAGNOSIS 6 2.8 mm, ovigerous 2 3.0 mm. Pleonite 5 dorsally smooth. 
Pleotelson with middorsal longitudinal rounded ridge bearing several 


Limnoria multipunctata 197 


Figure 87. Limnoria multipunctata: A, pleotelson; Limnoria pfefferi: B, pleotelson; 
Limnoria platycauda: C, pleotelson; Limnoria saseboensis: D, pleotelson; Limnoria 
simulata: E, pleotelson; Limnoria tuberculata: F, pleotelson. 


button-shaped tubercles in posterior half; posterior margin and lateral crests 


tuberculate. 


RECORDS Puerto Rico; Jamaica; Twin Cays, Belize. 
Japan; Kai Islands, South Pacific. 


198 FLABELLIFERA * LIMNORIIDAE 


Limnoria pfefferi Stebbing, 1904 
Figure 87B 


DIAGNOSIS d 3.8 mm, ovigerous ¢ 4.0 mm. Pleonite 5 with conspicuous 
middorsal Y-shaped carina. Pleotelson basally with pair of submedian 
rounded ridges; lateral crests lacking tubercles. 


RECORDS Florida Keys; Bahamas; Puerto Rico; U.S. Virgin Islands; Twin 
Cays and Man o’War Cay, Belize; Yucatan Peninsula, Mexico. 
Minikoi Atoll and Aldabra Atoll, Indian Ocean; Philippines; New Guinea; 


Panama. 


Limnoria platycauda Menzies, 1957 
Figure 87C 


DIAGNOSIS 6 2.5 mm, ovigerous 2 2.6 mm. Pleonite 5 with broad middor- 
sal longitudinal rounded ridge. Pleotelson lacking dorsal ornamentation; 
posterior margin and lateral crests bearing tubercles. 


RECORDS Cuba; Puerto Rico to Curacao; Cozumel, Mexico; Twin Cays 
and Man o’War Cay, Belize. 
Aldabra Atoll, Indian Ocean. 


Limnoria saseboensis Menzies, 1957 
Figure 87D 


DIAGNOSIS <6 3.5mm. Pleonite 5 with submedian pair of ridges, converging 
slightly posteriorly. Pleotelson basally with submedian pair of anteriorly tu- 
berculate ridges; posterior margin and lateral crests tuberculate. 


RECORDS Miami, Florida. 
Japan; Fiji. 


Limnoria simulata Menzies, 1957 
Figure 87E 


DIAGNOSIS 6 3.8 mm, ovigerous 2 4.0 mm. Pleonite 5 with obscure me- 
dian longitudinal groove. Pleotelson basally with submedian pair of tubercles 
and small lateral tubercles, latter often difficult to detect; lateral crests 
tuberculate. 


RECORDS Florida Keys; U.S. Virgin Islands; Gulf of Mexico. 


Paralimnoria andrewsi 199 


Limnoria tuberculata Sowinsky, 1884 
Figure 87F 


DIAGNOSIS <6 2.8 mm, ovigerous 2 3.0 mm. Pleonite 5 with two anterior 
tubercles, one middorsal posterior tubercle, area between tubercles de- 
pressed. Pleotelson basally with middorsal tubercle, followed by pair of sub- 
median tubercles, all three tubercles having short obscure carina; posterior 
margin and lateral crests tuberculate. 


RECORDS Rhode Island to Venezuela; Cuba; Man o’War Cay, Belize; Gulf 
of Mexico. 

Uruguay; West Africa; Mediterranean; Black Sea; India; Hong Kong; 
Hawaii; Australia; California. 


REMARKS This species has frequently been recorded under the name Lim- 
noria tripunctata Menzies, 195la. 


Limnoria unicornis Menzies, 1957 
Figure 88A,B 


DIAGNOSIS d 2.6 mm, ovigerous 2 2.6 mm. Mandibular palp of one arti- 
cle. Pleonite 5 with somewhat obscure Y-shaped ridge middorsally. 
Pleotelson in d with strong basal slightly curved middorsal tubercle; lateral 
crests lacking tubercles. 


RECORDS Bahamas; Man o’War Cay and Twin Cays, Belize. 
Caroline Islands; Palau; Society Islands. 


Paralimnoria Menzies, 1957 


DIAGNOSIS Antennular flagellum of five articles. Antennal flagellum of five 
or six articles. Mandibular incisor with rasp and file. Pleopod 5, rami bearing 
marginal setae. Uropodal rami subequal in length, each with clawlike apex. 


Paralimnoria andrewsi (Calman, 1910) 
Figure 88C,D 


DIAGNOSIS 6 2.6 mm, 2 2.6 mm. Pleonite 5 with or without triangular 
middorsal depressed area. Pleotelson with basal submedian pair of tubercles 
either obscurely or strongly carinate; lateral crest tubercles of variable 
strength. 


200 FLABELLIFERA * LIMNORIIDAE 


Figure 88. Limnoria unicornis: A, pleotelson, d; B, pleon, d, in lateral view. 
Paralimnoria andrewsi: C’, pleonite 5 and pleotelson; D, uropod. Phycolimnoria clarkae: 
E, pleonite 5 and pleotelson; F, uropod and pleotelson in lateral view. 


RECORDS Florida Keys; Puerto Rico; Twin Cays, Belize; Curacao. 
Christmas Islands, Indian Ocean; Samoa; Hawaii; Japan. 


REMARKS Menzies (1957) discusses three forms of this species: Forma typ- 
ica, which lacks a central depressed area dorsally on pleonite 5 and has a pair 
of submedian obscurely carinate tubercles on the pleotelson; Forma A, which 
has a triangular depressed area dorsally on pleonite 5 and a pair of subme- 


FLABELLIFERA * SEROLIDAE 201 


dian tubercles supported by strong carinae on the pleotelson; Forma B, hav- 
ing a triangular depressed area dorsally on pleonite 5 and an obscurely cari- 
nate pair of tubercles on the pleotelson. Given that at least two of these forms 
have been recorded occurring together, it would seem that this is merely a 
highly variable species. 


Phycolimnoria Menzies, 1957 


DIAGNOSIS Mandibular incisor lacking rasp and file. Uropodal rami une- 
qual, exopod longer than endopod, latter usually with clawlike apex. 


REMARKS Most species of Phycolimnoria are algal borers, frequently encoun- 
tered in the holdfasts of brown algae such as Macrocystis, Laminaria, and 
Sargassum. The one species recorded from the Caribbean, P. clarkae, however, 
has only been taken from decaying wood. 


Phycolimnoria clarkae Kensley and Schotte, 1987 
Figure 88E,F 


DIAGNOSIS <6 4.3 mm, ovigerous 2 3.3—4.4 mm. Uropodal exopod less 
than half length of endopod, straight, tipped with short squat claw. Pleonite 
5 with broad raised middorsal region having irregular bumps. Pleotelson 
wider than long, with two rounded submedian ridges basally, becoming ob- 
solete posteriorly. 


RECORDS Bahamas; Twin Cays, Belize. 
Aldabra Atoll, Indian Ocean. 


Family Serolidae Dana, 1852 


DIAGNOSIS Body dorsoventrally depressed. Eyes present or absent. 
Cephalon fused with pereonite 1 dorsally. Mandible bearing palp. Maxillipe- 
dal palp of one to four articles. Pereonites 2—4 with coxae demarked; per- 
eonites 5 and 6 with coxae not demarked; pereonite 7 narrow, lacking free 
lateral margins. Pereopod | in d and ¢ subchelate, pereopod 2 subchelate or 
ambulatory in 6, ambulatory in @. Pleonites 1 and 2 free, articulated, re- 
mainder of pleonites fused with telson. Pleopods 1—3 small, natatory; 
pleopods 4 and 5 large, operculate. Uropods lateral, biramous. 


REMARKS The serolids reach their greatest diversity (and their greatest size 
of up to 80 mm in length) in the southern oceans, with few species extending 


202 FLABELLIFERA * SPHAEROMATIDAE 


into the subtropics and tropics. The deep- and abyssal-dwelling species usu- 
ally lack eyes. The animals are epibenthic, living in the upper few centime- 
ters of the bottom sediment, where they are scavengers and carnivores. 


Serolis Leach, 1818 


DIAGNOSIS Body markedly dorsoventrally flattened. Coxal plates produced 
laterally. Mandible having lacinia mobilis and single spine. Maxillipedal 
palp of three articles (rarely two to four). Pereopod 2 exhibiting sexual di- 
morphism, subchelate in d, ambulatory in 2. Pleopods 1—3, peduncles elon- 
gate, rami subelliptical. Pleopod 3, exopod uniarticulate. 


Serolis mgrayi Menzies and Frankenberg, 1966 
Figure 89 


DIAGNOSIS 6 4.5 mm, ovigerous 2 4.7 mm. Eyes present. Cephalon with 
two middorsal tubercles. Pereonites 2—4 each with faint rounded tubercle 
just mesial to coxal suture. Pereon and pleon with faint middorsal longitudi- 
nal carina bearing small blunt tubercle on posterior margin of each segment. 
Pleonites 1 and 2 with lateral margins not contributing to body outline, over- 
lapped by pereonite 6. Pleotelson broadly triangular, with lateral carina in 
anterior half; apex truncate. Uropodal rami reaching to or slightly beyond 
pleotelsonic apex. 


RECORDS Off North Carolina, 18-34 m; off South Carolina, 22 m; off 
Georgia, 18—47 m; Florida Keys, 18-88 m; Trinidad; Venezuela, 95 m; Flor- 
ida, Gulf of Mexico, 11-88 m. 


Family Sphaeromatidae H. Milne Edwards, 1840 


DIAGNOSIS Antennular peduncle of three articles, antennal peduncle of five 
articles. Mandible stout, lacinia mobilis and molar usually well developed, 
palp of three articles. Maxillipedal palp of five articles. Mouthparts in some 
genera metamorphosed and somewhat reduced in ovigerous 2. Pleon of five 
partially or completely fused pleonites, often indicated by lateral sutures, 
plus dorsally convex and sometimes inflated pleotelson. Uropods lateral, ex- 
opod free if present, endopod fused with sympod. Sexual dimorphism often 
marked, especially in pleotelsonal structure. Animal often capable of con- 
globating or folding over. Young brooded in internal pouches or anterior or 
posterior pockets; oostegites variable in number, if present. 


FLABELLIFERA * SPHAEROMATIDAE 203 


Figure 89. Serolis mgrayi: A, 3; B, pereopod 1; C, pereopod 2, d. 


REMARKS Right into the 1980s this family was routinely divided into three 
groups, based on the structure of the two posterior pairs of pleopods: 
Platybranchiatae—pleopods 4 and 5 with both rami membranous and lack- 
ing branchial pleats; Hemibranchiatae—pleopods 4 and 5 with branchial 
pleats on endopods only; Eubranchiatae—pleopods 4 and 5 with branchial 
pleats on both rami. These three “groups” were recognized formally as sub- 
families by Hurley and Jansen (1977) but the names were not based on con- 


204 FLABELLIFERA * SPHAEROMATIDAE 


tained genera and were replaced with current subfamily names by Bowman 
(1981) and Iverson (1982), the latter providing diagnoses for all five sub- 
families. Four of these are represented in the Caribbean area; the fifth, the 
Tecticipitinae, contains only the single primarily Pacific genus Tecticeps. 
While the subfamilial status now appears to be resolved, many of the gen- 
era still require unambiguous diagnoses. The work of Harrison (1984) on the 
structure of the female broodpouch, with its various components of 
oostegites, internal pouches, and anterior and posterior pockets (Figure 90), 
along with the metamorphosis of the female mouthparts (see Figure 96) has 
helped enormously to standardize the genera. Nevertheless, these features of 
the female remain unknown in several genera. Further, with this stabilization 
based on females, many problems of incorrect generic designation have been 
uncovered. In this work, Harrison’s generic diagnoses are followed as far as 
possible. Where uncertainty exists, this is indicated. In some cases, we may 
still be unaware of existing problems: future work will without doubt result in 
the shifting of species to different genera, as well as in the creation of new 


genera. 


Key to subfamilies of Sphaeromatidae 


1. Pereopod 1 prehensile in both sexes; pereopod 2 prehensile only in ¢ 


SR o Be tiers MRNA aye elena SeaUd oe rene ck aeRO iy cichasy Pai a ascent Ancininae 

Pereopods: | and 2 ambulatory, si 5.2 den os fa es ee eee 2 

2. Pleopods 4 and 5 lacking branchial pleats ............. Cassidininae 

Pleopods 4 and 5 with branchial pleats on endopods ............... 3 

3. Pleopods 4 and 5 with branchial pleats on both rami ... Dynameninae 
Pleopods 4 and 5 with branchial pleats on endopods only 

Sess. ctaNe, Seta ia Gita ede on aN Ratha tine 0 reo. ie ge eae Sphaeromatinae 


Subfamily Ancininae Tattersall, 1905 


DIAGNOSIS Body markedly dorsoventrally depressed. Cephalon fused me- 
dially with pereonite 1. Pereopod 1 prehensile in d and 2. Pereopod 2 pre- 
hensile in 6 only. Pleopods 4 and 5 similar, lacking branchial pleats. 
Uropods uniramous. 


Ancinus belizensis 205 


anterior pocket 


oostegite 
posterior pocket 


internal pouch 


Figure 90. Diagrammatic representation of 2 sphaeromatid, showing marsupial 
structures (adapted from Harrison, 1984). 


Ancinus H. Milne Edwards, 1840 


DIAGNOSIS Eyes dorsal. 2 mouthparts not metamorphosed. Mandibular 
molar absent; palp of three articles. Maxilla | of single ramus, endite rudi- 
mentary. Maxilla 2 of two rami. with oostegites absent; brood held in two 
opposing pockets, opening as narrow ventral slit between pereopods 4. Pleon 
consisting of short anterior pleonite with free lateral margin, plus broadly 
triangular pleotelson. Pleopod | uniramous, endopod absent. Pleopod 2 op- 
erculiform. Pleopod 3, exopod of single article. Uropod lacking exopod, sym- 
pod not laterally expanded. 


Key to species of Ancinus 


1. Pleotelson as long as basal width, apex narrowly rounded .. . brasiliensis 
Pleotelson with basal width greater than length, apex subtruncate 
Bt ce OOo en ch soa Ba coca ila: alga int ke Ta a A One en belizensis 


Ancinus belizensis Kensley and Schotte, 1987 
Figure 91A—C 


DIAGNOSIS 6 4.1 mm, ? 2.8 mm. Body oval, about twice longer than wide. 
Dorsal integument strongly pitted. Antennular flagellum of 12 articles; an- 


206 FLABELLIFERA * SPHAEROMATIDAE 


Figure 91. Ancinus belizensis: A, 2; B, pereopod 1 3; C, pereopod 2 ¢. Ancinus 
braziliensis: D, adult (from Glynn and Glynn, 1974). 


tennal flagellum of 10 articles. d pereopod 2, dactylus strongly curved, 
reaching to proximal lobe of propodus. Pleopod 2 about 2.5 times longer than 
basal width. 


RECORDS Carlson Point, Belize, in seagrass flats, 0.5 m. 


Ancinus brasiliensis Lemos de Castro, 1959 
Figure 91D 


DIAGNOSIS 6 7.0mm, 2 6.0 mm. Body about twice longer than wide. Dor- 
sal integument smooth. Antennular flagellum of 17 articles; antennal 
flagellum of 10 articles. d pereopod 2, dactylus strongly curved, reaching to 
midlength of posterior margin of carpus. Pleopod 2 almost three times longer 
than basal width. 


Cassidinidea 207 


RECORDS Brazilian coast from Rio de Janeiro northward, 1.5 m; Costa 
Rica, Panama; shallow infratidal below sandy beaches. 


REMARKS Glynn and Glynn (1974) discussed color polymorphism in this 
species. 


Subfamily Cassidininae Iverson, 1982 


DIAGNOSIS Cephalon not medially fused with pereonite 1. Pereopod | am- 
bulatory. Pleopods 4 and 5, both rami lacking transverse pleats, outer rami 
unsegmented. Pleopod 5, outer ramus with low subapical squamiferous pro- 
tuberances. Pleotelsonic apex entire. Uropods with exopods reduced. 


REMARKS The genus Dies has twice been recorded from the Caribbean: D. 
arndti Ortiz and Lalana, 1980, from Cuba, and D. barnardi Carvacho, 1977, 
from Guadeloupe. This genus is distinguished from Cassidinidea solely on the 
basis of the penial structure: biramous in Cassidinidea, uniramous in Dies. 
Harrison (1984) has pointed out that the separation of these two genera has 
not been satisfactorily resolved. The penis of neither the Cuban nor the 
Guadeloupan species has been illustrated, but the whole-animal illustrations 
of both look suspiciously like Cassidinidea ovalis. Examination of material of D. 
barnardi from the Paris Museum supports the view that this species was based 
on immature material of C. ovalis. Neither of the so-called species of Dies are 
dealt with in this work, both being regarded as junior synonyms of C. ovalis. 


Key to genera of Cassidininae 


1. Frontal lamina visible dorsally between antennular bases; two basal 
articles of antennular peduncle not expanded .......... Cassinidinea 
Frontal lamina not visible between antennular bases; two basal articles 
of antennular peduncle broadly expanded ........ Paraleptosphaeroma 


Cassidinidea Hansen, 1905b 


DIAGNOSIS Body strongly dorsoventrally depressed. Eyes dorsal, situated 
at posterolateral corners of cephalon. Latter somewhat sunken into pereonite 
1. Frontal lamina expanded, visible dorsally between antennular bases. An- 
tenna directed laterally. Pleon consisting of one free pleonite having short 
free lateral margin, plus broadly triangular pleotelson. Uropodal endopod 


208 FLABELLIFERA * SPHAEROMATIDAE 


well developed, fused with sympod; exopod markedly reduced. Penial rami 
elongate, separate. 2 mouthparts not metamorphosed. Oostegites absent. 
Brood housed in pouch formed by opposing pockets overhanging ventrum, 
opening by slit between fourth pereopods. 


Key to species of Cassidinidea 


«Posterior mareginvef pleotelsoni truncate’ tic... use }. sees ae ovalis 
Postenior marginvef pleotelson rounded: . :.o21.2..24e sass. ap neer mosaica 


Cassidinidea mosaica Kensley and Schotte, 1987 
Figure 92A 


DIAGNOSIS. 6 1.8 mm, ovigerous 2 1.6 mm. Body twice longer than wide. 
Dorsal integument bearing close-packed flattened tubercles. Pleotelson tri- 
angular, with posterior margin narrowly rounded, dorsally convex, basally 
inflated. 


RECORDS Carrie Bow Cay, Belize, 1.5—10 m; in silty sand and rubble be- 
tween patch reefs and coral buttresses. 


Cassidinidea ovalis (Say, 1818) 
Figure 92B—E 


DIAGNOSIS 6 and 2 3.6 mm. Body width slightly less than half length. 
Dorsal integument smooth. Pleotelson with raised anteromesial area, but 
lacking sculpture; posterior margin truncate. 


RECORDS New Jersey to Florida, in marsh mud and among dead leaves, 0— 
1 m; Trinidad; Belize; Panama; Dominica; Louisiana and Vera Cruz, Gulf of 
Mexico. Known from waters of less than 1%o to 35%o. 


Paraleptosphaeroma Buss and Iverson, 1981 


DIAGNOSIS Body oval in outline, entire circumference with transparent 
flange of fused setae on two expanded basal articles of antennule, on per- 
eonites, pleonite 1, and uropods. Expanded basal articles of antennules con- 


Paraleptosphaeroma 209 


Figure 92. Cassidinidea mosaica: A, 3. Cassidinidea ovalis: B, 3; C, pereopod 1; D, 
pleopod 4; E, pleopod 5. Paraleptosphaeroma glynni: F, ¢. 


tiguous in midline. Single articulated pleonite with short free lateral margin. 


Uropodal sympod and endopod fused; exopod articulated, much shorter 
than fused endopod. 


210 FLABELLIFERA * SPHAEROMATIDAE 


Paraleptosphaeroma glynni Buss and Iverson, 1981 
Figure 92F 


DIAGNOSIS 6 2.58 mm, ovigerous ¢ 2.38 mm. Pleotelson basally broad, 
tapering to notched posterior margin. Fused uropodal endopod and sympod 
of each side almost touching posterior to pleotelsonic apex. 


RECORDS Portsmouth, Dominica, intertidal rock pools. 
Punta Paitilla, Pacific Panama. 


REMARKS Buss and Iverson (1981) demonstrated that this species displays 
sequential protogynous hermaphroditism, and that the change from female 
to male seems to be mediated by social conditions, especially the proportion 
of males to females. The principal food source for this species was shown to 
be abascan bryozoans. 


Key to genera of Dynameninae 


1, Pleotelson: very similar im ‘both sexes) 4. .c26.07 Gh oe yo ea 2 

Pleotelson showing marked sexual dimorphism ..................-. 3 

2. Cephalon and pleotelson smooth, lacking ridges .......... Ischyromene 

Pleotelson and cephalon with ridges: 2.3... 0h ee oe ce Cerceis 

3. JWrepods lamellar ‘both; sexes-4..4).\2)2.ta 10-4 pete seers <= or eee ee + 
Uropods lamellar in ¢, endopod reduced, exopod elongate-cylindrical 

UNS Geo ese 4B esaia: Des oe lado eS eal Na, oS Ree ee 5 


4. Ovigerous @ lacking oostegites; 6, pleopod 2 copulatory stylet basally 
broad, distally tapering, extending to or beyond ramus . . Dynamenella 
Ovigerous 2 with one pair of oostegites on pereonite 4; 6, pleopod 2 
copulatory stylet narrow, extending well beyond ramus ... . Paradella 


5. 6, strong median lobe in pleotelsonic notch reaching well beyond 


margin; ovigerous 2 with three pairs of oostegites ........ Discerceis 
3, pleotelsonic notch with short median lobe, if present; ovigerous 2 
with three or four, pairs, of costegites’ .7. 7... .: 0-4 eee 6 


6. Ovigerous ¢ with three pairs of oostegites; d pleotelsonic notch 
lacking “marginal teeth ‘or median lobes =.) <= ee ae Geocerceis 
Ovigerous ¢ with four pairs of oostegites; d pleotelsonic notch with 
marginal teeth and/or median lobe’..-% 002. see cet eee Paracerceis 


Discerceis lal 


Subfamily Dynameninae Bowman, 1981 


DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods | and 2 am- 
bulatory. Pleopods 4 and 5, both rami having branchial pleats. Pleopod 4, 
exopod unjointed, usually lacking setae, endopod with few setae at most. 
Pleotelsonic apex often with terminal notch or foramen, especially in d. 


Uropods biramous. 


Cerceis H. Milne Edwards, 1840 


DIAGNOSIS Mouthparts metamorphosed in ¢. Broodpouch of four pairs of 
oostegites on pereonites 1—4, overlapping in midline. Brood held in four pairs 
of internal pouches. Pockets absent. 


“Cerceis” carinata Glynn, 1970 
Figure 93A 


DIAGNOSIS 6 3.8 mm, @ 3.9 mm. Dorsal integument, especially posteri- 
orly, finely pustulose. Cephalon with three pairs of rounded dorsolateral 
carinae, one middorsal carina, not reaching posterior margin. Pereonite | 
with two ventrolateral carinae. Pleopod 3, exopod biarticulate. Pleotelson 
similar in d and @. Pleotelson with basal middorsal inflated area flanked by 
two smaller lateral swellings, with carina in midline almost reaching pos- 
terior margin; posterolateral margins converging to narrow, slightly concave 
posterior margin. d: Penes elongate, basally fused. Pleopod 2, copulatory 
stylet basally broad, tapering distally, articulating mediodistally on endo- 
pod; exopod with three enlarged distal plumose setae. 


RECORDS’ Venezuela, 5—7 m. 


REMARKS Several differences (in the male penial structure, copulatory 
stylet, antennular peduncle, and pleonal sutures) between Cerceis carinata and 
the definition of the genus (Harrison and Holdich, 1982; Harrison, 1984) 
indicate that this species has not been placed in the correct genus. Until fresh 
ovigerous females and mature males are available, the generic position must 
remain uncertain. 


Discerceis Richardson, 1905 


DIAGNOSIS Mouthparts in 2 metamorphosed. Broodpouch formed by four 
pairs of oostegites on pereonites 1—4, overlapping in midline. Brood held in 


212 FLABELLIFERA * SPHAEROMATIDAE 


Figure 93. “Cerceis” carinata: A, 3. Discerceis linguicauda: B, 3. Dynamenella 
acutitelson: C, 2; D, pleon 3. Dynamenella angulata: E, @. 


internal pouches (number unknown). Pockets absent. d with uropodal endo- 
pod and sympod fused, very short; exopod elongate, cylindrical. 


Dynamenella 213 


Discerceis linguicauda (Richardson, 1901) 
Figure 93B 


DIAGNOSIS 6 7.2 mm. Dorsal integument, especially of posterior half, with 
numerous scattered granular tubercles. Uropodal endopod and sympod 
fused, very short, exopod elongate, subcylindrical and slightly bowed. Ante- 
rior half of pleotelson inflated, with three elongate rounded ridges (each com- 
posed of two contiguous tubercles) ending posteriorly in subacute tubercle; 
posterior margin trilobed, median lobe broadly rounded, with subacute tu- 
bercle at base, lateral lobe truncate, well separated from median lobe. Head 
and pereonite | not fused. Frontal lamina visible dorsally between antennal 
bases. Penes short, separate. Copulatory stylet basally relatively broad, dis- 
tally broadly rounded. 


RECORDS Cape Catoche, Yucatan, Mexico, 48-50 m. 


REMARKS This species is known only from the four male syntypes. 


Dynamenella Hansen, 1905b 


DIAGNOSIS Species exhibiting obvious sexual dimorphism. Both sexes lack- 
ing processes on pereon and pleon. Uropodal rami lamellar. Exopod of 
pleopod 3 with or without articulation. 2: Mouthparts not metamorphosed. 
Broodpouch lacking oostegites, but formed by two opposing ventral pockets 
opening in midline between fourth pereopods. Apex of pleotelson with notch, 


Key to species of Dynamenella 


fers with pleotelsonic foramen) «)..2.. 0205.28.05 ene enn eee eee ees 2 
3 lacking foramen but with notch, or appearing entire; 2 pleotelson 
UALR AIOLEM VISMONC 2 dioica iis soe are see ss s,s ewes se acutitelson 


2. 3 with four strong pleotelsonic ridges; 2 with subcircular pleotelsonic 
fondimensiaen er iikimonise anes: bos cies Es bile ed ee quadrilirata 
6 lacking pleotelsonic ridges; ¢ with posterior margin of pleotelson 
Suite teas Ar Bhs RRS te ies ad eek Ecards perforata 


214 FLABELLIFERA *« SPHAEROMATIDAE 


groove, or foramen. d: Penes basally fused, rami long, tapering. Copulatory 
stylet proximally broad, tapering to acute tip, reaching to or just beyond 
apex of endopod. Uropods broader than in 2. Posterior pleotelson with dor- 
sally directed foramen connected to apex by narrow slit. 


REMARKS The species described by Richardson (1901) as Dynamene angulata 
from No Name Key, Florida, and referred to by some authors as a Dyna- 
menella, while figured here (Figure 93E), is not included in the present key. 
The species is known only from immature females; correct generic placement 
is thus not possible. 


Dynamenella acutitelson Menzies and Glynn, 1968 
Figure 93C,D 


DIAGNOSIS 6 3.5mm, ? 2.3 mm. d: Pereonites 4—6 with transverse ridge 
over dorsum, ridge interrupted to form short median section. Pleotelson with 
two submedian and two lateral rounded tubercles basally, two submedian, 
poorly defined ridges in central area; posterior margin tapering in dorsal 
view, with slit either just visible or appearing entire. In lateral view, posterior 
pleotelson seen to be laterally compressed, forming narrow groove. 


RECORDS Puerto Rico, intertidal rocks and algae. 


REMARKS Menzies and Glynn (1968) described this species with two vari- 
eties, the holotype as D. acutitelson var. typica, and 11 paratypes as D. acu- 
titelson var. glabrothorax. The major difference between these varieties lay in 
the presence of transverse ridges on pereonites 4—6 in ¢ypica and their absence 
in glabrothorax. The holotype, however, at 3.5 mm, would seem to be a mature 
male, while all the paratypes are smaller. The differences described by Men- 
zies and Glynn (1968) may thus be due to immaturity. As further compara- 
tive material is lacking, these varieties (or whatever their true status) are not 
recognized here. 

Menzies and Glynn (1968, fig. 30a) illustrate D. acutitelson var. glabrothorax 
as having scattered tiny granules over the dorsal integument. These were not 
seen when the type material was reexamined. 

Harrison and Holdich (1982) placed this species in Paradella, based on the 
literature. However, the penes for both varieties are shown as short and sepa- 
rate, as in Jschyromene. Again, until further mature males and ovigerous 
females are seen, the generic placement of this species must remain in doubt. 


Geocerceis barbarae DS 


Dynamenella perforata (Moore, 1901) 
Figure 94A,B 


DIAGNOSIS 6 3.2mm, 2 2.6 mm. d: Pleon bearing two low rounded sub- 
median “mounds.” Pleotelson with strongly convex anterior two-thirds, with 
T-shaped foramen. Pleon and pleotelson with numerous scattered small tu- 
bercles. Uropodal rami broadly ovate, outer margins crenulate. : 
Pleotelson broadly rounded in dorsal view, posterior margin entire. Inner 
uropodal ramus distally subacute. 


RECORDS Bermuda to Puerto Rico, intertidal coral rubble and algae, and 
under chiton Acanthopleura granulata; Dominican Republic; Cuba. 


Dynamenella quadrilirata Kensley, 1984 
Figure 94C—H 


DIAGNOSIS 6 2.6mm, 2 2.5 mm. 6: Two low rounded submedian tuber- 
cles on last pleonite. Anterior half of pleotelson inflated, with four rounded 
longitudinal ridges; posterior half tapered, somewhat dorsally flexed, with 
cordate foramen. Uropodal rami distally rounded, outer margins crenulate 
to dentate. 2: Lacking pleonal tubercles. Pleotelson inflated, unornamented, 
posterior margin forming subcircular foramen. 


RECORDS Carrie Bow Cay, and Twin Cays, Belize; intertidal to 3 m. 


Geocerceis Menzies and Glynn, 1968 


DIAGNOSIS Ovigerous 2 with mouthparts metamorphosed. Broodpouch 
with three pairs of oostegites, on pereonites 2—4, just overlapping in midline. 
Brood held in internal pouches (number unknown). Pockets absent. Uropo- 
dal rami lamellar, shorter than pleotelson. d uropodal endopod fused with 
sympod, very short; exopod elongate, club shaped. Pleopod 2 with copula- 
tory stylet articulating distally on endopod. 


Geocerceis barbarae Menzies and Glynn, 1968 
Figure 95A—C 


DIAGNOSIS 6 3.3mm, 2 2.5 mm. Pleopod 3 exopod of single article. Pleon 
with two elongate sutures reaching lateral pleon margin. d: Frontal lamina 
expanded into ventrally directed beaklike process. Penes separate, relatively 


216 FLABELLIFERA * SPHAEROMATIDAE 


Figure 94. Dynamenella perforata: A, 3; B, pleon 2. Dynamenella quadrilirata: C, 3; 
D, pleon ¢; E, pleon 2, lateral view; F, pleopod 2 3; G, pleopod 4; H, pleopod 5. 


elongate. Pleonite 5 with three dorsal tubercles near posterior margin. 
Pleotelson with raised anterocentral area having two lateral longitudinal 
rounded ridges; apex notched. 2: Uropodal exopod and endopod subequal, 
lamellar. Pleotelson as in ¢. Frontal lamina not produced as in d. 


RECORDS Puerto Rico, intertidal to 3 m, in coral rubble. 


Ischyromene aT 


Figure 95. Geocerceis barbarae: A, 3; B, pleon 2; C, 6, lateral view. Ischyromene 
barnardi: D, o. 


Ischyromene Racovitza, 1908 


DIAGNOSIS mouthparts not metamorphosed. Broodpouch of three pairs 
of oostegites on pereonites 2—4, overlapping in midline. Large posterior 
pocket covering posterior ventrum, opening anteriorly between fourth per- 
eopods. Brood housed in ventral body wall. Sexual dimorphism not pro- 


218 FLABELLIFERA * SPHAEROMATIDAE 


nounced. Uropodal rami lamellar. d pleopod 2 with copulatory stylet 
basally narrow, reaching to or just beyond distal margin of endopod. 


Ischyromene barnardi (Menzies and Glynn, 1968) 
Figure 95D 


DIAGNOSIS 6 4.5mm, ¢ 3.7 mm. Both sexes lacking processes on pereon 
and pleon. Accessory unguis of pereopods often bifid. Pleopod 3, exopod of 
single article. Uropodal rami lamellar. 6: Pereonite 7, posterior margin 
bilobed. Penes short, separate to base. 


RECORDS Puerto Rico, intertidal. 


Paracerceis Hansen, 1905b 


DIAGNOSIS Pleopod 3 exopod with transverse suture in distal half. Pleon 
with two long sutures reaching to posterolateral margin. 6: Penial rami 
short, separate. Pleotelson with basal area strongly vaulted; deep posterior 
notch sometimes having denticles on inner margins, and/or median tooth at 
base of notch. Uropodal endopod short, fused with sympod; exopod elongate, 
club shaped. ¢: Mouthparts metamorphosed. Mandible fused with 
cephalon. Broodpouch of four pairs of oostegites, three posterior pairs over- 
lapping. Brood retained in internal pouches. Uropodal rami subequal, lamel- 
lar. Pleon usually less ornamented than in 6, and with shallower median 
notch lacking teeth. 


Key to species of Paracerceis (P. nuttingi not included) 


1. d, pleotelsonic notch narrow, with median basal tooth; 2, pleotelson 
dorsally. unormmamented: 5 «fic canis idare nto ee ee ee ee 2 

3, pleotelsonic notch wide, with lateral teeth; 2, pleotelson dorsally 
with: tubercles: 2.5.<25te vod.c Sal sicie oie elas Se eee 3 


2. 3, median tooth of pleotelsonic notch almost as long as notch; 
subacute median tubercle on anterior pleotelson; 2, pleotelson with 
posterior margin faintly concave, not notched .............. edithae 

3, median tooth less than half length of notch; pleotelson with blunt 
rounded median tubercle; 2, pleotelson with distinct posterior notch 


88 DUE A Sie GEO a Se ee ee glynni 


Paracerceis caudata 219 


3. 3, pleotelsonic notch deep, margins usually with two teeth on each 
side; strong median tubercle on anterior pleotelson bluntly bifid; @, 
pleotelson with one or two rounded median tubercles and 2 smaller 
BUI RMCS EON Gd CI USTCMED Nyko) sete ire lais con dhe oan doesn 2 egahees caudata 

3, pleotelsonic notch shallow, with tiny lateral denticles; median 
tubercle of pleotelson conical, acute; °, pleotelson with three large 
conical acute tubercles and several smaller scattered tubercles in 
CANA MACHTMEN AIL on tus eles ok con recM Maes oe so sees aks SACP See Ee cohenae 


Paracerceis caudata (Say, 1818) 
Figure 96 


DIAGNOSIS. 6 8.1 mm, 2 6.4 mm. ¢: Pleotelson with blunt median bifid 
tubercle, with two smaller tubercles on each side. Pleotelsonic notch usually 
with two strong denticles on each margin, basal median tooth lacking. 
Uropodal exopod reaching well beyond pleotelson, slightly bowed, with 2—4 
setose bumps on outer margin. 2: Pleonite 5 with three low tubercles. 
Pleotelsonic apex broadly rounded in dorsal view, with two rounded median 
tubercles and two smaller tubercles on each side. Uropodal rami subequal, 
lamellar, outer distal angle of each acute. 


RECORDS’ Bermuda; New Jersey to Florida Keys; Yucatan to Venezuela; 
Turks and Caicos Islands; Cuba; Puerto Rico; Bahamas; Jamaica; Haiti; St. 
Maartens, 0.2—127 m; St. Lucia; Gulf of Mexico. Found in the following 
algae: Caulerpa, Halimeda, Turbinaria, Amphiroa, Laurencia, Dictyota; between 
sponges and tunicates on red mangrove roots; in coral rubble; in spur and 
groove zone of reefs, lagoon, back reef, seagrass flats, and fringing 
mangroves. 


REMARKS Menzies and Glynn (1968:55, fig. 22f) named and figured P. 
caudata var. brevipes from Puerto Rico. This variant was characterized as hav- 
ing the margins of the pleotelsonic notch lacking denticles. Given the con- 
siderable variation in ornamentation in this species, we feel that no validity 
can be given to the name “brevipes.” 

This is the commonest sphaeromatid in the Caribbean, and it has very 
broad ecological requirements, being found in a wide range of habitats and 
depths. 


220 


FLABELLIFERA * SPHAEROMATIDAE 


Ni 


Al IME Ay, ly 


Figure 96. Paracerceis caudata: A, 3; B, pleon 2; C, mandible 6; D, maxilla 1 6; E, 
maxilla 2 ¢d; F, maxilliped d; G, mandible 2; H, maxilla 1 2; J, maxilla 1 2; J, 


maxilliped °. 


Paracerceis cohenae Kensley, 1984 

Figure 97A,B 
DIAGNOSIS 6 10.0 mm, 2 7.9mm. d: Pereonites each with median tuber- 
cle and several smaller lateral tubercles near posterior margin of somite. 
Pleonite 5 with large median conical tubercle. Anterior two-thirds of 


Paracerceis glynni 221 


pleotelson inflated, faintly tripartite, with strong median conical tubercle; 
notch in posterior margin shallow, with low median tooth and tiny lateral 
denticles; posterolateral margins finely dentate. Uropodal exopod cylindri- 
cal, distally denticulate, six to seven times longer than basal width. @: Per- 
eon and pleon much as in 4, but pleotelsonic notch shallower and 
posterolateral margins not denticulate. Uropodal rami subequal, lamellar, 
exopod with distolateral angle acute. 


RECORDS Carrie Bow Cay, Belize, 15-16 m. Only known from sponge 
Callispongia plicifera growing on outer reef slope. 


Paracerceis edithae Boone, 1930 
Figure 97C—E 


DIAGNOSIS 6 4.0 mm, 2 3.1 mm. d: Posterior three pereonites and 
pleonites each with irregular row of small tubercles near posterior margin, 
densely setulose tubercles becoming spinose more posteriorly. Pleotelson 
with strong median conical tooth in anterior half, flanked by convex spinose 
mound. Pleotelsonic notch deep, with elongate median basal tooth bearing 
strong acute tooth at its base. Lobes of posterior pleotelsonic margin broad, 
flattened, margins denticulate. Uropodal exopod tuberculate, tapering, api- 
cally acute. 2: Integument much less tuberculate-spinose than in d. Imma- 
ture 2, posterior margin of pleotelson with faintly rounded median lobe. In 
mature @, posterior margin distinctly trilobed. Uropodal rami subequal, 
lamellar, distally rounded, with tiny distolateral spine on exopod. 


RECORDS Bahamas, 60—66 m, in vase sponge; Haiti; Puerto Rico, 20-25 m. 


Paracerceis glynni Kensley, 1984 
Figure 97F,G 


DIAGNOSIS 6 6.4mm, 2 5.2 mm. d: Integument becoming strongly setose 
and tuberculate posteriorly from about pereonite 5. Posterior margin of inf- 
lated anterior area of pleotelson bearing strong median conical tubercle and 
smaller acute lateral tubercle, with low swelling beneath each lateral tuber- 
cle. Posterior notch deep, narrow, with small basal median tooth, lobes form- 
ing notch tricuspid, outer cusps recurved dorsally. Uropodal exopod fairly 
straight, cylindrical, apically acute. 2: Body far less setose and tuberculate 
than d. Pleotelson with strongly inflated anterior area having very faint mid- 
dorsal tubercle; notch well marked, formed by triangular lobes of 


222 FLABELLIFERA * SPHAEROMATIDAE 


Figure 97. Paracerceis cohenae: A, 3; B, pleotelson, 2. Paracerceis edithae: C, 
pleotelson, 2; D, mature 6; E, immature 6. Paracerceis glynni: F, 3; G, pleotelson, 


2. Paracerceis nuttingi: H, &. 


pleotelsonic margin. Uropodal rami subequal, flattened, endopod with distal 
margin faintly trituberculate; exopod with few distal tubercles. 


RECORDS Alligator Light, Florida, 11 m; Carrie Bow Cay, Belize, 11—15.2 


Paradella DS 


m, from green alga Halimeda sp. on forereef, and from sponge Aphysina 
fistularis. 


Paracerceis nuttingi (Boone, 1921) 
Figure 97H 


RECORDS Barbados; Puerto Rico, 1.5 m, from Cymodocea seagrass, and coral 
rubble and sponges. 


REMARKS The types of this species from Barbados consist only of females 
(total length 4.1 mm). Menzies and Glynn (1968) record an immature male 
from Puerto Rico with an incipient pleotelsonic notch. This specimen, how- 
ever, still has the subequal lamellar uropodal rami. The mature male, with 
the characteristically reduced uropodal endopod and cylindrical exopod, is 
unknown. The possibility exists that this is not a true Paracerceis. 


Paradella Harrison and Holdich, 1982 


DIAGNOSIS Both sexes lacking processes on pereon and pleon. Marked sex- 
ual dimorphism. Accessory unguis of pereopods simple, not bifid. Pleopod 3 


Key to species of Paradella 


1. Pereonite 7 with projecting bilobed flange; pleon and pleotelson finely 
Dita distincelygoramulate os 203 3. aac tie can: aes es OR oes So OM Ohl eas Za 
PICGUN AGN PI EOLe ISON SIMOOUA. 2 2c ac. <n Spskus (rn ee Re, a Ss. - 3 


2. 3d with pleotelsonic foramen distinctly heart shaped, with median 
point; four submedian tubercles of pleotelson in d and 2 somewhat 
elongate; 2 pleotelson posteriorly narrowed, slit visible dorsally 

3 with pleotelsonic foramen wider than long, but lacking median 
point; four submedian tubercles of pleotelson in d small, rounded, 
obscure in 2; @ pleotelson posteriorly truncate, slit not visible 
(SPOT gS | | Pee ae UE OE ge ea ORR: Aieee 1 27 Sapo eta plicatura 


3. Tubercles on pleotelson in d and ¢ small to obscure; d with 
plcotelsomic foramen subcircular 2.665.055.0605 00s oan quadripunctata 
Tubercles on pleotelson broadly rounded mounds in d and 2; 6 with 
pleotelsonic foramen wider than long ................. tumidicauda 


224 FLABELLIFERA * SPHAEROMATIDAE 


exopod with articulation. Uropodal rami lamellar. 2: Mouthparts not meta- 
morphosed. One pair of oostegites arising from pereonite 4, short, not reach- 
ing midline. Brood held in pouch formed by two opposing pockets covering 
entire ventrum, opening by transverse slit between 4th pereopods. d: Penes 
long, basally fused. Copulatory stylet basally narrow, extending further be- 
yond endopod than in Dynamenella. Uropods broader than in &. 


Paradella dianae (Menzies, 1962b) 
Figure 98A—C 


DIAGNOSIS 6 3.4mm, ovigerous 2 3.7 mm. d: Pereonite 7, posterior mar- 
gin broadly bilobed. Pleonite 5 with two rounded submedian tubercles. 
Pleotelsonic foramen distinctly heart shaped with median point. Pleotelson 
with four submedian and two lateral tubercles in d and ¢, plus median 
tubercle at base of foramen in d; tubercles tending to be elongate and sub- 
carinate. Uropodal rami lamellar, margins finely crenulate, relatively 
broader than in @. @: Pleotelsonic slit wide, dorsally visible. 


RECORDS Key West, Florida; Puerto Rico, intertidal. 
Baja California, intertidal. 


Paradella plicatura (Glynn, 1970) 
Figure 98D,E 


DIAGNOSIS 6 4.1 mm, ¢ 3.6 mm. Pleon and pleotelson with tiny scattered 
tubercles. 6: Pereonites 5—7 with posteriorly directed flanges, that of per- 
eonite 7 largest, bilobed. Pleotelson with four submedian and two lateral 
discreet rounded tubercles. Pleotelsonic foramen wider than long, with basal 
bulge but lacking median point. 2: Pleotelson with wide posterior slit not 
visible dorsally, posterior margin appearing truncate. Pleotelsonic tubercles 
less marked than in 6. 


RECORDS Jamaica, under red mangroves; Margarita Island, Venezuela, 
shallow infratidal. 


Paradella quadripunctata (Menzies and Glynn, 1968) 
Figure 98F,G 


DIAGNOSIS 6 2.5mm, 2 2.5 mm. Pleonite 5 with two low rounded subme- 
dian tubercles. Pleotelson with four low submedian rounded tubercles and 


Paradella quadripunctata 225 


Figure 98. Paradella dianae: A, 3; B, pleopod 2 3; C, pleon 2. Paradella plicatura: 
D, 3; E, pleon 2. Paradella quadripunctata: F, 3; G, pleon 2. Paradella tumidicauda: 
H, pleon @ (from Glynn, 1970); J, 6. 


four smaller lateral tubercles. ¢: Pleotelsonic foramen subcircular, ventral 
margins of foramenal tube barely touching. : Pleotelson posteriorly nar- 
rowly tapered, slit becoming tubelike, dorsally visible; four submedian tuber- 
cles less marked than in 6. 


RECORDS Bermuda; Florida; Dominican Republic; Puerto Rico, intertidal; 
U.S. Virgin Islands, intertidal to 1 m. 


226 FLABELLIFERA * SPHAEROMATIDAE 


Paradella tumidicauda (Glynn, 1970) 
Figure 98H,I 


DIAGNOSIS 6 6.7 mm, 2 6.5 mm. <d: Last pleonite with two submedian 
swellings. Pleotelson with four submedian broadly rounded swellinglike tu- 
bercles and two pairs of lateral tubercles. Foramen wider than long, posterior 
contiguous borders of foramen each bearing rounded swelling. 2: Pleotelson 
with four submedian swellinglike tubercles, sometimes with two obscure lat- 
eral tubercles; posterior slit not visible dorsally, area surrounding slit 
swollen, horseshoe shaped. 


RECORDS Margarita Island, Venezuela, from among intertidal barnacles. 


Subfamily Sphaeromatinae H. Milne Edwards, 1840 


DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods 1 and 2 am- 
bulatory. Pleopods 4 and 5, endopods having branchial pleats, exopods un- 
pleated, membranous, of two articles. Uropods biramous. 


Key to genera of Sphaeromatinae 
1.’ Uropodal<exopod with outer margin serrate ... 25. se o2 Sphaeroma 
Uropodal exopod with outer margin entire or faintly crenulate ...... 2 


2. 3, pleotelsonic notch with median lobe; 2, pleotelsonic apex barely 
notched with rounded median lobe; 2, mouthparts metamorphosed 


alate Mate CIS ee NE EOE MIA SO Loe eee at eer ay eee 3 

Pleotelson entire to very faintly notched in d and 2; 2, mouthparts 
not metamorphosed; with three pairs of oostegites ..... Exosphaeroma 

3. Mature 6, uropodal exopod about twice length of endopod; 2 with 
three pairs OhOOSteEsItEs :. soca c ee tee nae ween eae Harrieta 

3, uropodal rami subequal or exopod shorter than endopod; ? with 
{Our Pairs OL GOSTEPITES \...5-5.8 uc step oasian ce cee sortase Cymodoce 


Cymodoce Leach, 1814 


DIAGNOSIS Pleon with two elongate straight parallel incomplete sutures on 
each side. Pleotelsonic apex with marked notch bearing median tooth. 


Cymodoce ruetzleri PT, 


Pleopod 5, exopod of two articles, distal article with apex and internal mar- 
gin covered with fine teeth, anterior surface with long distally toothed boss; 
proximal article with two small toothed bosses at internodistal angle. d: 
Maxillipedal palp articles 2—4 bearing setigerous lobes. Penial rami elongate, 
separate. Pleon usually more tuberculate than in 2. Uropodal exopod lamel- 
lar, shorter than endopod. 2: Mouthparts metamorphosed. Broodpouch 
formed by four pairs of oostegites arising from pereonites 1—4, overlapping in 
midline. Brood housed in five pairs of internal pouches. 


“Cymodoce” barrerae (Boone, 1918) 
Figure 99A,B 


DIAGNOSIS 2 7.5mm. 2: Body dorsally strongly vaulted, unornamented. 
Frontal lamina distally broadly rounded, lateral shoulders rounded. 
Mouthparts not metamorphosed. Pleotelson anteriorly strongly inflated with 
barest indication of two submedian swellings; posterior margin trilobed, with 
median lobe strong, narrowly rounded, outer lobes much smaller and ventral 
to median lobe. Uropodal endopod distally obliquely truncate; exopod dis- 
tally acute. 


RECORDS Cabanas, Cuba. 


REMARKS This species is known only from the nonovigerous female 
holotype. Loyola e Silva (1960) placed the species in Cymodoce, based on a 
female specimen from Brazil. As the mouthparts are not metamorphosed, 
this does not agree with the present concept of Cymodoce, but with neither 
ovigerous females nor males available, the correct generic placement cannot 
be determined. 


Cymodoce ruetzleri Kensley, 1984 
Figure 99C—G 


DIAGNOSIS d¢ 5.0 mm, 2 4.2 mm. ¢: Integument with numerous small 
tubercles, becoming densely setose posteriorly. Pleonite 4, posterior margin 
broadly bilobed. Pleotelson bearing pair of strong conical tubercles with 
acute tips, each tubercle flanked by low rounded tubercle; apex trilobed, 
outer lobes triangular, acute, sharp spine at base of incision, median lobe 
apically blunt. Uropodal exopod apically acute, oval in cross section, endo- 
pod and sympod fused, somewhat flattened, apex triangular with strong 
tooth. 2: Pleotelson with two conical apically acute tubercles, apex barely 
notched, with short rounded lobe slightly offset from posterior margin. Both 


228 FLABELLIFERA * SPHAEROMATIDAE 


Figure 99. “Cymodoce” barrerae: A, 2; B, frontal lamina. Cymodoce ruetzleri: C, 3; D, 
pleon in lateral view, 6; E, pleon @; F, pleopod 4; G, pleopod 5. 


uropodal rami flattened; exopod with tiny apical tooth, endopod distally 
truncate-rounded, with small mediodistal tooth. 


RECORDS Carrie Bow Cay, Belize, 0.5—13 m; in algal clumps, reef crest 
rubble, and seagrass flats. 


Exosphaeroma alba 229 


Exosphaeroma Stebbing, 1900 


DIAGNOSIS Maxillipedal palp articles 2—4 produced medially into lobes. 
Pereonites 6 and 7 dorsally unarmed. Pleopod 3, exopod biarticulate. d: 
Penes short, separate. Copulatory stylet of pleopod 2 elongate, slender. 
Pleotelson lacking strong apical notch. 2: Mouthparts not metamorphosed. 
Broodpouch of three pairs of oostegites on pereonites 2—4; oostegites short, 
not reaching midline. Brood held in four pairs of internal pouches. 


Key to species of Exosphaeroma 
1. Pleotelson with posterior margin entire, evenly convex ............. 2 
Pleotelson with posterior margin faintly notched or trilobed ........ 3 


2. Frontal lamina with length less than 1.5 times greatest width diminuta 
Frontal lamina with length almost two times greatest width 


See oe tea aeRO STING Me cc ane hn oP ake terest productatelson 

3. Pleotelson with posterior margin faintly tnlobed, and with three low 
rounaged, tubercles anteriorly... 2. h.5. slosh sx teers soo yucatanum 
Pleotelson with posterior margin faintly notched .................. 4 


4. Pleotelson posteriorly broadly notched; two rounded submedian 
tubercles;on' inflated midregion® = 2:..0. 2.062 22 Os Peed antillense 
Pleotelson with faint narrow notch posteriorly; lacking dorsal tubercles 


OU A Sa 2 > RR a ER ORS oe A ME 7a ES OS LOU EE CS.» =. ee ROE alba 


Exosphaeroma alba Menzies and Glynn, 1968 
Figure 100A—C 


DIAGNOSIS 6 2.0 mm, 2 2.3 mm. Frontal lamina anteriorly broadly 
rounded, basally slightly wider than midlength. Pleotelson similar in d and 
2; anterodorsally inflated and unornamented, posteriorly tapering to slight 
median notch, seen in dorsal view. Uropodal rami distally shallowly serrate, 
exopod 2.5 times longer than wide. 


RECORDS Puerto Rico, intertidal to 0.5 m; in algae on rocks, and under 
Chiton tuberculatus and C. marmoratus. 


230 FLABELLIFERA * SPHAEROMATIDAE 


Figure 100. Exosphaeroma alba: A; B, frontal lamina; C, uropod. Exosphaeroma 
antillense: D; E, frontal lamina; F, uropod. Exosphaeroma diminuta: G; H, frontal 
lamina; J, uropod. Exosphaeroma productatelson: J; K, frontal lamina; L, uropod. 
Exosphaeroma yucatanum: M, pleon (from Richardson, 1905). 


Exosphaeroma antillense Richardson, 1912d 
Figure 100D,F 


DIAGNOSIS @ 5.0 mm. Frontal lamina anteriorly tapering to subacute apex. 
Pleotelson with two broadly subconical submedian tubercles on inflated an- 


Exosphaeroma yucatanum 231 


terior area; posterior margin subtruncate to very faintly emarginate. Uropo- 
dal exopod distally crenulate, length slightly more than twice greatest width; 
endopod with faint distal notch. 


RECORDS Montego Bay, Jamaica. 


REMARKS The single ovigerous female holotype is the only known specimen 
of this species. The overlapping oostegites suggest that this may not be an 
Exosphaeroma. 


Exosphaeroma diminuta Menzies and Frankenberg, 1966 
Figure 100G—I 


DIAGNOSIS <6 2.2 mm. Frontal lamina widest at midlength, anteriorly 
truncate-rounded. Pleotelson with posterior margin broadly rounded. 
Uropodal rami not quite reaching pleotelsonic apex; exopod margin distally 
crenulate. 


RECORDS Chesapeake Bay to Florida; Venezuela; sand dwelling, intertidal 
and shallow subtidal. 


Exosphaeroma productatelson Menzies and Glynn, 1968 
Figure 100J—L 


DIAGNOSIS 6 2.5mm, ¢ 1.5 mm. Sexes essentially similar. Frontal lamina 
widest at midlength, where slight shoulder apparent, anteriorly broadly 
rounded, 1.6 times longer than wide. Pleotelson unornamented, anteriorly 
inflated, posterior margin entire, evenly convex. Uropodal exopod distally 
shallowly serrate, almost four times longer than wide; endopod wider than 
exopod. Broad lateral patches of pigment on pleotelson in both sexes. 


RECORDS Puerto Rico, intertidal to 0.5 m, in algae on rocks; Texas, Gulf of 
Mexico. 


Exosphaeroma yucatanum (Richardson, 1901) 
Figure 100M 


DIAGNOSIS. Frontal lamina anteriorly tapering from widest point to sub- 
acute apex, proximally narrower than at midlength. Pleotelson posteriorly 
obscurely trilobed, median lobe narrowly rounded, longest; three low 
rounded tubercles on pleotelson in anterior region. 


232 FLABELLIFERA * SPHAEROMATIDAE 


RECORDS Cape Catoche, Yucatan, Mexico, 48 m. 


REMARKS This species was described from a single specimen which has 
since been lost. The true generic placement of this species is thus undeter- 
mined and full description awaits the finding of more material. 


Harrieta Kensley, 1987c 


DIAGNOSIS ¢ with mouthparts metamorphosed. Broodpouch of three pairs 
of oostegites on pereonites 2—4, overlapping in midline; brood held in five 
pairs of internal pouches. Uropodal rami subequal, lamellar in 2, exopod 
twice length of endopod and oval in cross section in 6. Pleopod 2 in 6 with 
copulatory stylet articulating basally on endopod, curved, barely reaching 
apex of endopod. Penes basally fused, rami slender, elongate, tapering. 


Harrieta faxoni (Richardson, 1905) 
Figure 101A,B 


DIAGNOSIS 6 6.0 mm, ¢ 6.5 mm. G: Frontal lamina with broad slightly 
convex anterior margin. T'wo low rounded submedian tubercles on cephalon 
near posterior margin. Two rounded submedian tubercles on last pleonite. 
Pleotelson anteriorly inflated with two submedian tubercles; posterior mar- 
gin trilobed. ¢: Essentially similar to d, but posterior margin of pleotelson 
less markedly trilobed, with median lobe longer, and uropodal rami subequal 
in length. 


RECORDS Florida to Texas, Gulf of Mexico, intertidal and subtidal in 
Thalassia, Halodule, and Syringodium seagrass beds, in salinities of 7%o to 36%o. 


Sphaeroma Bosc, 1802 


DIAGNOSIS Maxillipedal palp with three distal articles poorly developed, 
lacking lobes; fringe of robust plumose setae with swollen bases on internal 
margin of endite; distal margin of endite with simple setae. Pereopods 1—3 
with plumose setae on ischium and merus. Posterior margin of pleotelson 
entire, similar in d and 2. Pleopod 3, exopod uniarticulate. Uropodal ex- 
opod with outer margin serrate. Able to conglobate. ¢d: Penes short, 
rounded, separate. Pleopod 2, copulatory stylet articulating basally on endo- 
pod, slender, reaching well beyond rami. 2: Mouthparts not meta- 


Figure 101. Harrieta faxoni: A, 3 ; B, pleon 2. C, Sphaeroma quadridentata; D, 
Sphaeroma terebrans; E, Sphaeroma walkeri. 


234 FLABELLIFERA * SPHAEROMATIDAE 


morphosed. Three pairs of overlapping oostegites arising from pereonites 2— 
4 (but S. terebrans has anterior pair rudimentary). 


REMARKS ‘The genus Sphaeroma is one of the few sphaeromatids in which the 
number of oostegites varies, from the diagnostic three pairs, through two 
normal pairs (as in S. terebrans), to having the oostegites completely absent 
(as in S. annandalei). 

Jacobs (1987) has provided a useful reevaluation of the European, Medi- 
terranean, and northwest African species of Sphaeroma and related genera. 


Key to species of Sphaeroma 


1. Pleotelson posteriorly bluntly triangular, with 4 strong anterior 


tubercles 
Fo MS MSS Ae bos He ed re mae ecto rent eaatate ete nate, aaa ca eS terebrans 
Pleotelson: posteriorly broadly rounded .).2- 5a o5-. coe ie oe 2 
2. Pleotelson dorsally smooth or with few low tubercles ..... quadridentata 
Pleotelson dorsally with numerous strong tubercules .......... walkeri 


Sphaeroma quadridentata Say, 1818 
Figure 101C 


DIAGNOSIS 6 11.0 mm, ¢ 8.0 mm. Pleotelson anteriorly inflated, some- 
times with few low rounded tubercles, posteriorly flattened to concave; pos- 
terior margin entire, broadly rounded. 


RECORDS New England to Florida; Gulf of Mexico, intertidal to 1 m, often 
in pilings and partially submerged dead tree trunks, and commonly associ- 
ated with barnacles. 


Sphaeroma terebrans Bate, 1866 
Figure 101D 


DIAGNOSIS. 6 10.0 mm, @ 11.5 mm. Pereonite 7 with pair of submedian 
and pair of lateral tubercles. Dorsal pleon densely tuberculate. Posterior 
pleonite with pair of submedian acute tubercles. Pleotelson anteriorly with 
submedian pair and lateral pair of tubercles, posteriorly rounded-triangular. 


FLABELLIFERA * TRIDENTELLIDAE 235 


RECORDS Virginia to Florida; Belize; Cuba; Venezuela to Brazil; Gulf of 
Mexico. 

Nigeria, east coast of southern Africa, India, Sri Lanka, Thailand, Indo- 
nesia, Philippines, Australia. 


REMARKS There is no agreement on whether this species is synonymous 
with §. destructor Richardson, 1897. This latter (if distinct) bores into wood 
pilings in estuarine waters, while S. terebrans is found in the prop roots of the 
red mangrove tree, Rhizophora mangle. In this habitat, the isopods are inter- 
preted either as being destructive agents (e.g., Rehm and Humm, 1973) or as 
promoting increased root growth (Simberloff et al., 1978). It is unlikely that 
the bored wood itself is a source of food for the isopods; rather, as with the 
genus Limnoria, the food is probably detritus or fungi and bacteria growing on 
the wood fragments in the burrows or on the setae of the appendages. 


Sphaeroma walkeri Stebbing, 1905 
Figure 101E 


DIAGNOSIS 6 9.5mm, 2? 10.0 mm. Pereonites 3—7 with transverse row of 
large rounded tubercles. Last pleonite with row of prominent tubercles and 
smaller scattered tubercles laterally. Pleotelson anteriorly inflated, posteri- 
orly concave and cuplike, with four irregular longitudinal rows of large tu- 
bercles plus many small scattered tubercles. Posterior margin rounded, en- 
tire to irregularly crenulate. Uropodal endopod with several rounded 
tubercles on dorsal surface; exopod with row of smaller tubercles on ventral 
surface. 


RECORDS Probably pan-tropical. Florida to Puerto Rico, intertidal. 


Family Tridentellidae Bruce, 1984 


DIAGNOSIS Eyes well developed. Pereonites 2—7 with distinct coxae. Pleon 
consisting of five free pleonites plus pleotelson. Mandible with acute incisor; 
lacinia mobilis absent; molar present; palp of three articles. Maxilla 1, outer 
ramus styliform with three to five strong terminal spines, and several short 
recurved subapical spines. Maxilla 2 uniramous, biarticulate, bearing small 
sometimes tridentate spines or scales distally. Maxillipedal palp of five arti- 
cles; endite slender, lamellar, usually lacking coupling hooks. 


236 FLABELLIFERA * TRIDENTELLIDAE 


Tridentella Richardson, 1905 


DIAGNOSIS Body dorsally often bearing spines, tubercles, or carinae, more 
developed in ¢ than in 2. Frontal lamina narrow, pentagonal. Antennular 
peduncle of three articles; antennal peduncle of five articles. Mandibular 
molar weakly sclerotized. Pereopods 1—3 weakly prehensile; pereopods 4—7 
ambulatory. Copulatory stylet of pleopod 2 rodlike, arising proximally on 
mesial margin of endopod. Pleopod 5 endopod lacking marginal setae. 


REMARKS Delaney and Brusca (1985) provide useful taxonomic and dis- 
tributional comments on the family Tridentellidae. 


Tridentella virginiana (Richardson, 1900b) 
Figure 102 


DIAGNOSIS 6 9.5 mm, ovigerous $ 9.5—11.0 mm. Cephalon and pereon 
dorsally smooth, pleon minutely granular. Uropodal rami with distal mar- 
gins faintly dentate, apically narrowly rounded, endopod wider and slightly 
longer than exopod. Pleotelson basally wider than middorsal length; pos- 
terior margin broadly rounded to subtruncate. 


RECORDS Nova Scotia to Florida; off Georgia, 550 m; Gulf Stream off Key 
West, 220 m. 


Suborder Gnathiidea Leach, 1814 


DIAGNOSIS Eyes usually well developed, rarely on short lateral processes, 
occasionally absent. Mandibles in ¢ greatly enlarged, projecting anteriorly 
from cephalon, not used in feeding. Mandibles lacking in 2. Mouthparts of 
praniza larva styliform, with acute mandibles projecting anteriorly (see Fig- 
ure 103D). Pereopod 1 modified, forming second pair of broad opercular 
maxillipeds covering mouthparts, referred to as pylopods. Pereopods 2—6 
ambulatory. Pereonite 7 reduced, lacking pereopod. Pleonites separate, nar- 
rower than pereon. Uropods lateral, rami lamellar, forming tailfan with 
telson. Praniza larva with pereonites 4—6 enlarged, sometimes inflated. 
with pereonites 4—6 greatly inflated, forming broodpouch for internally 
brooded eggs (see Figure 103E). 


REMARKS The gnathiideans are entirely marine, most described species 
being from shallow waters. The males and females are frequently found in 
association with sponges and do not feed. The praniza larva is an efficient 
swimmer and has been recorded from shallow-water plankton, but is more 


GNATHIIDEA 237 


Figure 102. Tridentella virginiana: A, 2; B, pereopod 1; C, maxilla 1; D, mandible; 
E, maxilliped; F, maxilla 2. 


frequently encountered as a fish parasite, the favored site for sucking the 
host’s blood being in the nares. Upton (1987a, 1987b) has shed light on the 
unusual life history of at least one gnathiid genus, Paragnathia. 

The taxonomy of the Gnathiidae is based almost entirely on males, the 
praniza and females of most species being remarkably similar. 


238 GNATHIIDEA * GNATHIIDAE 


Family Gnathiidae Harger, 1879 


DIAGNOSIS’ As for the suborder Gnathiidea. 


Gnathia Leach, 1814 


DIAGNOSIS In addition to features mentioned in diagnosis of suborder: Eyes 
present in most species. Pylopod with two small articles distal to broad oper- 
cular article 2, terminal article minute. 


Key to species of Gnathia (d only) 


1. Anterior margin of cephalon with medial process or slightly convex .. 2 
Anterior margin of cephalon concave or lacking medial process ..... 9 


2. Anterior margin of cephalon broadly triangular, projecting, with small 


lateral’ -teethi tr, 202 aig oe Bae ee Te, ee ee triospathiona 
Anterior margin of cephalon not triangular and projecting .......... 3 

3. Cephalon and two free anterior pereonites dorsally granular ........ 4 
Cephalon and two free anterior pereonites smooth ..............--- a 


4. Lobe of outer margin of mandible notched; pereonite 5 twice wider 


thanumiddorsal lengthy 26.2024 2s ots es oes tos sae ee care velosa 

Lobe of outer margin of mandible rounded; pereonite 5 1.5 times wider 

thanmiddorsallength On 2 snc... role ots Se ene oo ae 6 

5. Anterior margin of cephalon with distinct medial process ..... virginalis 
Anterior margin of cephalon barely convex, lacking medial process 

SSE Ate SR tO Se Ny Ein? Cee eS rathi 

6. Inner proximal lobe of mandible distinct ................5---00: 7 

Inner proximal lobe of mandible indistinct ............--- samariensis 

7. Inner proximal lobe of mandible entire ..............+++--+s220+:: 8 
Inner proximal lobe of mandible with rounded toothlike marginal 

SEMUECLUROS. |) 50 Arta. 2 aie coy encire a eps heen ann eee se oer Kee og ren aaa johanna 

$.!Pereonites.3—5 poorly defined 2.6). ).c ovata weine : poe puertoricensis 

Pereonites.3—5 clearly detined rai Aisi, bas aise Sake ee magdalenensis 

9. Anterior margin of cephalon concave, lacking projections ..... gonzalezi 


Anterior margin of cephalon with four projections .......... beethoveni 


Gnathia magdalenensis 239 


Gnathia beethovenit Paul and Menzies, 1971 
Figure 103A 


DIAGNOSIS. 6 3.0 mm. Anterior margin of cephalon with two low tubercles 
flanking shallow medial notch plus slightly larger pair of lateral tubercles. 
Cephalon lacking dorsal tubercles. Pereonite 5 1.5 times wider than middor- 
sal length. Uropodal endopods reaching beyond telsonic apex. 


RECORDS Off Venezuela, 95 m. Colombia. 


Gnathia gonzalezi Miller, 1988 
Figure 103B 


DIAGNOSIS. ¢ 2.0 mm. Body smooth. Anterior margin of cephalon concave. 
Pereonites 3—5 distinct; pereonite 5 2.5 times wider than middorsal length. 
Cutting margin of mandible with four or five low rounded teeth. 


RECORDS Colombia, 30 m. 


Gnathia johanna Monod, 1926 
Figure 103C 


DIAGNOSIS 6 2.1 mm. Anterior margin of cephalon medially convex be- 
tween pair of submedian tubercles. Pereonites 4 and 5 poorly separated. 
Proximomedial lobe of mandible having four or five rounded crenulations, 
with tiny seta between adjacent crenulations. 


RECORDS’ U.S. Virgin Islands, 29-46 m; Colombia. 


Gnathia magdalenensis Miller, 1988 
Figure 103D 


DIAGNOSIS 6 3.0 mm. Anterior margin of cephalon with three tubercles, 
median tubercle slightly shorter than submedian pair. Cephalon with few 
scattered low granulations dorsally. Pereonite 5 about 1.5 times wider than 
middorsal length. Proximomedial lobe of mandible entire. 


RECORDS Carrie Bow Cay, Belize, intertidal; Colombia, 18 m. 


Figure 103. Gnathia beethhoveni: A, 6. Gnathia gonzalezi: B, 5 (after Miller, 1988). 
Gnathia johanna: C, 3. Gnathia magdalenensis: D, 3 (after Miiller, 1988). Gnathia 
puertoricensis: E, 3; F, praniza larva; G, ovigerous °. 


Gnathia triospathiona 241 


Gnathia puertoricensis Menzies and Glynn, 1968 
Figure 103E—G 


DIAGNOSIS <6 3.0 mm, ovigerous 2 1.8 mm. Anterior margin of cephalon 
having three tubercles between mandibular bases, medial tubercle narrower 
than submedian pair. Dorsal integument finely granular, with coarser gran- 
ules mediodorsal to eye. Pereonites 4 and 5 indistinctly separated. Mandible 
lacking proximomedial lobe. 


RECORDS Carrie Bow Cay, Belize, intertidal to 2 m; Puerto Rico, intertidal; 
Cuba. 


Gnathia rathi Kensley, 1984 
Figure 104A 


DIAGNOSIS 6 1.9 mm, ovigerous 2 2.2 mm. Frontal margin faintly convex 
to straight between single low lateral tubercle mesial to mandibular bases. 
Dorsal integument of cephalon and anterior two free pereonites coarsely 
granular. Lateral margins of telson faintly denticulate. Pereonites 4 and 5 
poorly separated. 


RECORDS Carrie Bow Cay, Belize, 1-36 m. 


Gnathia samariensis Miller, 1988 
Figure 104B 


DIAGNOSIS 6 2.0 mm. Anterior margin of cephalon with three tubercles, 
median tubercle slightly shorter than submedian pair; dorsal integument 
smooth. Pereonites 4 and 5 well differentiated; pereonite 5 about 2.2 times 
wider than middorsal length. Mandible lacking proximomedial lobe. 


RECORDS Colombia. 


Gnathia triospathiona Boone, 1918 
Figure 104C 


DIAGNOSIS <6 8.8 mm. Anterior margin of cephalon with broad-based tri- 
angular projection bearing three low teeth; deep V-shaped depression pos- 
terior to anterior margin, with low flanking granulations. 


RECORDS Off Key West, in Gulf Stream, 218 m. 


Figure 104. Gnathia rathi: A, 6; Gnathia samariensis: B, 3 (after Miller, 1988); 
Gnathia triospathiona: C, 3; Gnathia virginalis: D, 3; Gnathia velosa: E, 3 (after 
Miiller, 1988). 


MICROCERBERIDEA * MIGROCERBERIDAE 243 


Gnathia velosa Miller, 1988 
Figure 104E 


DIAGNOSIS. ¢ 1.5 mm. Anterior margin of cephalon with three tubercles, 
median tubercles slightly shorter and narrower than submedian pair. Dorsal 
integument of cephalon and anterior three pereonites granular. Pereonite 5 
about 2.5 times wider than middorsal length. Lateral lobe of mandible 
» notched. 


RECORDS Colombia. 


Gnathia virginalis Monod, 1926 
Figure 104D 


DIAGNOSIS. 6 2.2 mm. Anterior margin of cephalon with three tubercles, 
median tubercles slightly longer than submedian pair. Dorsal integument of 
cephalon and anterior three pereonites granular. Pereonite 5 about 1.7 times 
wider than middorsal length. Lateral lobe of mandible rounded. 


RECORDS U.S. Virgin Islands, 29 m; Colombia. 


Suborder Microcerberidea Lang, 1961 


DIAGNOSIS Cephalon free. Mandibles with reduced palp, or lacking palp. 
Maxillipedal palp of five articles. Pereon of seven free segments. Pereopod | 
subchelate; pereopods 2—7 ambulatory. Pleon of two free pleonites plus 
pleotelson. Pleopod | in d usually absent. Pleopod 2 modified for copulation. 
Pleopod 3 uniramous, opercular. Pleopod 4 biramous. Pleopod 5 reduced. 
Uropods usually uniramous or biramous. 


Family Microcerberidae Karaman, 1933b 


DIAGNOSIS’ Eyes absent. Body elongate, slender. Antennular peduncle of 
three articles; antennal peduncle of six to eight articles. Mandibular palp of 
single article; molar reduced to single stout fringed spine. Maxilla 2 reduced 
to single ramus bearing two distal fringed lobes. Pereopods 2—7 ambulatory, 
dactyli biunguiculate. 


REMARKS The species of the Microcerberidae are all very small (less than 2 
mm total length) and are most often found in interstitial habitats. They have 
been recorded from marine, brackish, and freshwater environments. 


244 MICROCERBERIDEA ¢ MICROCERBERIDAE 


The microcerberideans were often classified with the Anthuridea, mainly 
because of the similarity in body shape. Wagele (1983) however, has con- 
vincingly demonstrated the asellotan affinities of the group. 


Key to genera of Microcerberidae 


1. Maxillipedal palp articles 2 and 3 enlarged; basis of pereopods lacking 
SPUMOUS: PLOCESS! s.5. 542 clases eee sto ans tare) wean Sad Yvesia 
Maxillipedal palp articles 2 and 3 not markedly enlarged; basis of 
pereopads: with spinous Process, fe. fees bs ee eee Microcerberus 


Microcerberus Karaman, 1933b 


DIAGNOSIS Maxillipedal palp articles 2 and 3 not enlarged. Articles 2 and 3 
of antennal peduncle with spinous process. Basis of pereopods with spinous 
process. Propodus of pereopod 2 with two denticulate proximal spines. 


Microcerberus syrticus Kensley, 1984 
Figure 105A—E 


DIAGNOSIS. 6 1.1 mm, ¢ 1.1 mm. Tergal lobes of pereonites 2—4 rounded. 
Apical lobe of d pleopod 2 acute. 


RECORDS Carrie Bow Cay, Belize, interstitial in intertidal sand bar. 


REMARKS In addition to M. syrticus, six species of Microcerberus have been 
recorded from the Caribbean area: M. littoralis Chappuis and Delamare De- 
boutteville, 1956, from the Bahamas; M. minutus Coineau and Botosaneanu, 
1973, from Cuba; M. mirabilis Chappuis and Delamare Deboutteville, 1956, 
from the Bahamas; M. nunezi Coineau and Botosaneanu, 1973, from Cuba; 
M. renaudi Chappuis and Delamare Deboutteville, 1956, from the Bahamas; 
M. simplex Coineau and Botosaneanu, 1973, from Cuba. The reader is re- 
ferred to the original descriptions for separation of the species. 


Figure 105. Microcerberus syrticus: A, 6; B, pereopod 1; C, maxilliped; D, pereopod 
2; E, pleopod 2 6. Yuvesia striata (from Coineau and Botosaneanu, 1973): F, 
maxilliped; G, pereopod 1; H, pereopod 2. 


246 MICROCERBERIDEA * ONISCIDEA 


Yvesia Coineau and Botosaneanu, 1973 


Yvesia striata Coineau and Botosaneanu, 1973 
Figure 105F—H 


DIAGNOSIS ¢ 1.6 mm. Antennal peduncular articles 2 and 3 smooth, lack- 
ing spinous processes. Maxillipedal palp articles 2 and 3 enlarged. Bases of 
pereopods unarmed, lacking spinous processes. Propodus of pereopod | with 
single smooth proximal spine. Body having longitudinal ventrolateral striae. 


RECORDS Oriente, Cuba, interstitial on beach. 


Suborder Oniscidea Latreille, 1803 


DIAGNOSIS Compound eyes usually present. Antennules usually very short. 
Antennae with 4- or 5-articulate peduncle; flagellum varying from few arti- 
cles to multiarticulate. Mandibular palp present. Distal articles of maxillipe- 
dal palp often reduced. Coxae of pereonites 1—7 usually distinct, expanded. 
Pleopods respiratory, often with pseudotrachea; d with pleopod 2, and 
sometimes pleopod | as well, modified for copulation. Uropods terminal or 
subterminal with terete rami, or ventral and opercular, with reduced rami. 


REMARKS The Oniscidea includes all the isopods that have successfully in- 
vaded the terrestrial environment. While still in some degree reliant on exter- 
nal moisture, their morphological and behavioral adaptations have allowed 
them to live in almost all terrestrial habitats, from hot, dry deserts, through 
tropical rainforests and grasslands, to cold-temperate niches. Several forms 
have successfully inveigled themselves into termite or ant colonies, where 
with varying degrees of morphological adaptations they take advantage of 
the security of these habitats. A small number of species have evolved to live 
in more constantly wet habitats. Several species may be found in the marine 
intertidal, either living in and under piles of decomposing litter along the 
high-tide line, digging into beach sand, or sheltering in the damp cracks and 
crevices of rocky shores. A few may also be found in mangrove swamps. 

A breakdown of families, genera, and species is not provided for this sub- 
order, but those few species that are commonly encountered in intertidal 
habitats are dealt with individually. Schultz (1974, 1984) records several 
oniscidean isopods from the Caribbean area. 


ONISCIDEA 247 


Key to genera and species of littoral Oniscidea 


1. At least one uropodal ramus reaching well beyond outline of body ... 5 
Uropodal rami very short, not reaching beyond outline of body ..... 2 
2. Uropods ventral, not ‘visible in dorsal view .....'2.5...... 250s. 3 (Tylos) 
Mropeds visible in dorsal view, 25...2% oo... 2. 2 Armadilloniscus ninae 
3. Ventral extensions of pleonite 5 meeting in midline ....... Tylos niveus 
Ventral extensions of pleonite 5 not meeting in midline ............ 4 
4. Ventral extensions of pleonite 5 very short, obsolete ..... Tylos wegeneri 
Ventral extensions of pleonite 5 well separated ........ Tylos marcuzzii 


Ventral extensions of pleonite 5 just falling short of meeting in midline 
JP: Seep Sa I Oe nO ee PY APR CIRC 2 CAR mee Rana Tylos latreillei 


2. Uropodal rami both clongate, subequal .... 206. 66.6020. 6 (Ligia) 
Wropedal ram very unequal m length 22.2 2. dag ds ieee nt a ade oe 8 
6. Propodus of d pereopod | with distal rounded lobe ...... Ligia exotica 
Propodus of 6 pereopod | lacking rounded lobe .................. ‘| 
imaAmexval i mleopod 2 club shaped 252.2... 4-0... S24 ee Ligia olfersii 
mpEXor oc pleapod.? (bile. 7.2. fh. ge shes wads «Te dds Ligia baudiniana 
eoeantennal fagelum of two articles 254. o¢2626:...--¢ Rhyscotus texensis 
Antennal flagellum of three articles 2.00.5 cos. 5.5252 he- 9 (Vandeloscia) 


9. Endopod of ¢ pleopod | with large scalelike subapical process 
Ne er eRe href ec eto Ree ds he A Vandeloscia riedli 

Endopod of d pleopod | with small scalelike subapical process 
BN da ig eka), 5) 2s g Pg ac sess eo) Vandeloscia culebrae 


Armadilloniscus Ul yanin, 1875 


Armadilloniscus ninae Schultz, 1984 
Figure 106A 


DIAGNOSIS. 6 3.2mm, 2 4.1 mm. Uropodal sympod expanded to form part 
of body outline; rami set mesial to expanded base, with exopod half length of 
endopod. 


RECORDS Ambergris Cay, Belize; under damp objects along beach drift 
line. 


Figure 106. A, Armadilloniscus ninae. Ligia baudiniana: B; C, 3 pleopod 2 endopod. 
Ligia exotica: D, dactylus and propodus of pereopod 1; E, 6 pleopod 2 endopod. 
Ligia olfersii: F, 3 pleopod 2 endopod. G, Rhyscotus texensis. Tylos latreillei: H, 
ventral pleon. Tylos marcuzzi: I, ventral pleon (from Schultz, 1984). Tylos niveus: J, 
lateral view; K, ventral pleon. Tylos wegeneri: L, ventral pleon. Vandeloscia culebrae: 
M, apex of pleopod | endopod. N, Vandeloscia riedli. 


Rhyscotus texensis 249 


Ligia Fabricius, 1798 
Ligia baudiniana H. Milne Edwards, 1840 
Figure 106B,C 


DIAGNOSIS. 6 and 2 up to 22 mm. Antennal flagellum elongate, multiar- 
ticulate. Apex of d pleopod 2 bifid, with lateral lobe longer and more slender 
than mesial lobe. Uropods inserted terminally on pleotelson; sympods 
elongate-cylindrical; rami slender, elongate, subequal. 


RECORDS Bermuda; Bahamas; U.S. Virgin Islands; Antigua; Carrie Bow 
Cay, Belize; Bonaire; Aruba; Trinidad; Tobago; Gulf of Mexico. 


REMARKS As is typical in the genus Liga, this species may be seen on rocks 
and sea walls, as well as piles of drift debris at low tide. When disturbed, they 
run rapidly, to shelter in damp crevices and hollows. 


Ligia exotica Roux, 1828 
Figure 106D,E 


DIAGNOSIS. 6 28.5 mm, ovigerous 2 32.0 mm. Propodus of d pereopod | 
with rounded lobe on inner distal surface. Apex of d pleopod 2 club shaped, 


convoluted. 


RECORDS New Jersey to Uruguay; Indo-Pacific. 


Ligia olfersii Brandt, 1833 
Figure 106F 


DIAGNOSIS 6 20.0 mm, ovigerous 2 24.0 mm. Apex of d pleopod 2 simple, 
club shaped. 


RECORDS South Florida to Rio de Janeiro, Brazil; Texas, Gulf of Mexico. 


Rhyscotus Budde-Lund, 1885 
Rhyscotus texensis (Richardson, 1905) 
Figure 106G 


DIAGNOSIS. ¢ and 2 6.0 mm. Antennal flagellum of two unequal articles. 
Uropodal endopod at least twice length of exopod, inserted distally on base, 
exopod inserted distally on base. Pleotelson broadly triangular. 


RECORDS Carrie Bow Cay, Belize; Texas, Gulf of Mexico. 


250 ONISCIDEA 


Tylos Latreille, 1826 
Tylos latreillei Audouin, 1826 
Figure 106H 


DIAGNOSIS 6 12.8 mm, @ 13.0 mm. Ventral extensions of pleonite 5 not 
meeting in midline. 


RECORDS’ Bermuda; Cuba; Puerto Rico; Honduras. 
Mediterranean. 


Tylos marcuzzu Soika, 1954 
Figure 106] 


DIAGNOSIS. 6 6.6 mm. Antennal flagellum of four articles. Ventral exten- 
sions of pleonite 5 well separated. 


RECORDS Florida Keys; Bahamas; Leeward Islands; Ambergris Cay, Be- 
lize; under debris on sand beach drift line. 


Tylos niveus Budde-Lund, 1885 
Figure 106J,K 


DIAGNOSIS. 6 11.0 mm., 2 12.0 mm. Antennal flagellum of four articles. 
_ Ventral extensions of pleonite 5 expanded, medially contiguous. 


RECORDS Bahamas; Florida Keys; Cuba; Dominica; Lesser Antilles; Bo- 
naire; Curagao, under piles of decaying mangrove leaves at beach drift line; 
Carrie Bow Cay, Ambergris Cay, Belize, under deep piles of dead plant ma- 
terial on beach drift line; Tobago; Panama. 

Rio de Janeiro, Brazil. 


Tylos wegeneri Vandel, 1952 
Figure 106L 


DIAGNOSIS 6 10.5 mm, 2 15 mm. Antennal flagellum of three articles. 
Ventral extensions of pleonites short or nearly absent. Pleonite 5 lacking free 
lateral margins. 


RECORDS ‘Tobago; Venezuela, under decaying beach debris on drift line; 
Trinidad. 


VALVIFERA 251 


Vandeloscia Roman, 1977 
Vandeloscia culebrae (Moore, 1901) 
Figure 106M 


DIAGNOSIS 6 5.0 mm, 2 6.1 mm. Tiny lateral tubercles present on per- 
eonites. Endopod of pleopod | in d with small scalelike subapical process on 
laterally folded tip. 


RECORDS Florida Keys; U.S. Virgin Islands; Puerto Rico; under decaying 
plant material, especially Thalassia testudinea accumulated along beach drift 


line. 


Vandeloscia riedli (Strouhal, 1966) 
Figure 106N 


DIAGNOSIS 6 5.9 mm, ¢ 6.0 mm. Tiny obsolete tubercles present on all 
pereonites. Endopod of d pleopod 1 with large scalelike subapical process on 
laterally folded tip. 


RECORDS Yucatan Peninsula, Mexico; Ambergris Cay, Belize; Barbuda; 
Venezuela; Brazil. 

Gulf of Aqaba; Red Sea; northeastern coast of Africa; Madagascar; Bay of 
Bengal; St. Helena Is. 


Suborder Valvifera Sars, 1882 


DIAGNOSIS Pereopodal coxae, in addition to usual dorsal coxal plates, ex- 
panded ventrally to form plates. Penes situated ventrally on articulation be- 
tween pereon and pleon, or on pleonite 1. Pleonites and pleotelson variously 
fused. Uropods forming operculum covering over pleopods. 


Key to families of Valvifera 


1. Body often geniculate, flexed between pereonites 4 and 5; anterior 
pereopods setose for feeding, posterior pereopods ambulatory 
OTE Se A, ne ee One a ee ee Arcturidae 
Body never geniculate; all pereopods ambulatory .......... Idoteidae p 755 


252 VALVIFERA * ARCTURIDAE 


REMARKS Of the six families in the suborder, only two have been recorded 
in the Caribbean area, the Idoteidae and the Arcturidae. 


Family Arcturidae Sars, 1897 


DIAGNOSIS Pereonite | either distinct, or completely or imcompletely fused 
with cephalon. Anterior four pairs of pereopods directed anteriorly, usually 
strongly setose; posterior three pairs of pereopods ambulatory, used for cling- 
ing to substrate. Body often bent between pereonites 4 and 5. Uropods usu- 
ally biramous, with minute endopod concealed by larger exopod. Pleonites 
variously fused with pleotelson. Sexual dimorphism often marked. 


REMARKS Menzies and Kruczynski (1983) described three species of 
arcturids from the west coast of Florida, in depths of 55-73 m: Arcturella 
spinata, Arcturella bispinata, and Edwinjoycea horologium. These species are not 
covered here. 


Key to genera of Arcturidae 


1. Pereonite 1 not fused with cephalon; at least one free pleonite 
Jen Sales obo dks ain a aden ee St Pe oe a he Ao Lae) MES Thermarcturus 
Pereonite 1 fused with cephalon; pleonites fused with pleotelson 
SN eR EAN SE Bae wa Blk Oma in Me a aati Aa ii acne desk RP Astacilla 


Astacilla Cordiner, 1793 


DIAGNOSIS. Antennae at least half length of body. Pereopod 1 with strong 
terminal claw on dactylus. Pereopods 2—4 lacking dactyli. Endopod of 


Key to species of Astacilla 


1. Body integument lacking ornamentation .................. cymodocea 
Body, integument swith; spines or, tubercles..9. 656 citecye sears “sles 2 


2. Pereonite 4 in d and 2 with strong middorsal tubercle; pairs of spines 

lacking: On pereonitEs 280... seen oa eon ae oe eee regina 
Pereonite 4 lacking strong middorsal tubercle; pairs of spines on all 

pereonites °./) (G0 cu Vita nn seer teraes oe ne kre ete eee eas lasallae 


Astacilla regina 253 


pleopod 1 d with median notch and three specialized setae; pleopod 2 copu- 
latory stylet apically trifid. Pereonite 4 considerably longer than preceeding 
or following pereonite. 


Astacilla cymodocea Menzies and Glynn, 1968 
Figure 107A,B 


DIAGNOSIS. 6 6.4 mm, ovigerous ¢ 9.0 mm. Body cylindrical, ovigerous ? 
with pereonite 4 somewhat bulged, d with pereonite 4 elongate-cylindrical. 
Shallow groove marking fusion between cephalon and pereonite |. Pleonites 
fused with pleotelson, with two incomplete shallow dorsal grooves marking 
lines of fusion anteriorly. Pleotelson lacking any shoulders, posteriorly tape- 
red to narrowly rounded apex. 


RECORDS Florida Keys; Puerto Rico, 1.5 m, on Cymodocea sp. seagrass; Car- 
rie Bow Cay, Belize, 1-2 m, on Syringodium filiforme seagrass. 


REMARKS In life, A. cymodocea is bright green, blending in with its preferred 
substrate of seagrasses. 


Astacilla lasallae Paul and Menzies, 1971 
Figure 107C 


DIAGNOSIS 3.5 mm. Cephalon with large rounded area bearing pair of 
spines; all pereonites and two anterior fused pleonites bearing pair of short 
submedian spines. Pleotelson with strong anterior shoulder, posteriorly tri- 
angular, tapering sharply to narrowly rounded apex. 


RECORD Off Venezuela, 95 m. 


REMARKS This species is known only from the small female holotype, and 
until a mature male and ovigerous female are found, it cannot be confidently 
diagnosed. 


Astacilla regina Kensley, 1984 
Figure 107D—G 


DIAGNOSIS 6 6.5 mm, ovigerous 2 7.1 mm. Body strongly tuberculate, 
many tubercles acute. Cephalon with two submedian pairs of acute tuber- 
cles; fused pereonite 1 and pereonites 2 and 3 each with single middorsal 
acute tubercle. Pereonite 4 with strong middorsal tubercle situated in ante- 


254 VALVIFERA * ARCTURIDAE 


Figure 107. Astacilla cymodocea: A, 3; B, °. Astacilla lasallae: C, 3. Astacilla regina: 
D, 6; E, 2; F, pereopod 4; G, pereopod 1. Thermarcturus venezuelensis: H, 2 (from 
Paul and Menzies, 1971). 


rior half. Pleotelson with strong lateral shoulder in anterior half, second 
shoulder in posterior half, then tapering to rounded apex. 


RECORDS Carrie Bow Cay, Belize, on forereef slope, 27-36 m; Barbados, 
100-400 m; St. Lucia, 2—3 m, associated with crinoids. 


VALVIFERA * IDOTEIDAE 255 


Thermarcturus Paul and Menzies, 1971 


DIAGNOSIS Pereonite | not fused with cephalon. Pereonite 4 subequal in 
length to pereonite 3, not markedly elongate. Pereopods 2—4 having dactyli 
but lacking elongate setae. Body cylindrical, flexed between pereonites 4 and 
5. Pleon consisting of two free pleonites plus pleotelson. 


Thermarcturus venezuelensis Paul and Menzies, 1971 
Figure 107H 


DIAGNOSIS 2 4.5 mm. Cephalon, all pereonites, and anterior two pleonites 
each with submedian pair of dorsal tubercles, those on pereonites 2 and 3 
broad and expanded. Pleonite 2 with pair of bulbous lateral swellings, pos- 
terior margin triangular. Pleotelson with lateral shoulder anteriorly, posteri- 
orly triangular. 


RECORDS Off Venezuela, 95 m. 


REMARKS Only the holotype (which seems to be lost) is known of this spe- 
cies. Considerable uncertainty exists regarding some of the features. 


Family Idoteidae Fabricius, 1798 
Subfamily Idoteinae Dana, 1852 


DIAGNOSIS’ Flagellum of antenna either multiarticulate; clavate, i.e., with 
large basal articles and with or without one to four reduced distal articles; or 


Key to genera of Idoteinae 


antennal Mapenunr mulliariculate: ic. ..6 . 65's cede fan Idotea 
Antennal flagetlumiclavate es 2... lees ol eee oh Dae Stic 2S 2 
2. Pereopod 4 reduced, considerably smaller than pereopods 3 or 5 .... 3 


Pereopod 4 not reduced, of similar size to pereopods 3 and 5 
sss atk daha Riel APRESS A, WOR OER, OP ER dst Erichsonella 


3. Pleon consisting of three complete and one incomplete pleonites plus 
TENTS Rel TCT CARRS gh Sea Ee en SE a MMC ene Cleantioides 
Pleon consisting of two complete and two incomplete pleonites plus 
pleotelsion. @ cj aganst So hiins deen lees ler Beds at Miratidotea 


256 VALVIFERA ¢ IDOTEIDAE 


vestigial. Maxillipedal palp consisting of five or fewer articles. Uropods uni- 
ramous or biramous, rami usually much smaller than sympod. Pleonites 
variously fused with pleotelson; number of fused pleonites often indicated by 
lateral sutures or furrows. 


REMARKS Brusca (1984) has reviewed the phylogeny, evolution, and bio- 
geography of the subfamily Idoteinae, the only one of the five subfamilies 
recorded from the Caribbean. 


Cleantioides Kensley and Kaufman, 1978 


DIAGNOSIS Antennal flagellum a single clavate article. Maxillipedal palp of 
four or five articles. Pereopod 4 somewhat reduced. Uropod uniramous. 
Pleon consisting of three complete and one incomplete pleonites plus 
pleotelson. 


Cleantioides planicauda (Benedict, 1899) 
Figure 108A 


DIAGNOSIS Ovigerous ? 5.5 mm. Body parallel sided. Maxillipedal palp of 
five articles. Pleotelson posteriorly broadly rounded, with obliquely truncate 
subcircular dorsal area in posterior half. 


RECORDS Maryland to Florida; Puerto Rico; Panama; Louisiana, Gulf of 
Mexico, intertidal to 44 m; often in hollow stems and roots of seagrasses, and 
tubes of the polychaete Diopatra cuprea. 

Oaxaca, Pacific Mexico. 


REMARKS Cleantioides planicauda has been recorded only once in the eastern 
Pacific, where it occurs with the more common C. occidentalis (Richardson, 
1899). 


Key to species of Erichsonella 


Ll. .Pereonites with dotsal' spines q. 2o45 Sh chee ee fo Lh ceca soutien ae 2 
Pereonitesdacking dorsal ‘Spies! We ule eS ele ite es ssi scan attenuata 
2. Pereonites 1—4 with middorsal and lateral spines ............ floridana 


Pereonites 1—4 with middorsal spines only .................. filiformis 


Erichsonella 257 


Figure 108. A, Cleantioides planicauda; B, Erichsonella attenuata 3; C, Erichsonella 
filiformis 3; D, Erichsonella floridana 2; E, Idotea balthica 3; F, Idotea metallica 2; G, 
Miratidotea bruscai 2. 


Erichsonella Richardson, 1901 


DIAGNOSIS Antennal flagellum clavate. Maxillipedal palp of four articles. 
Uropod uniramous. Pleonites completely fused with pleotelson. 


REMARKS Pires (1984) reviewed the genus Erichsonella and did not recog- 
nize the subspecies E. filiformis tropicalis Menzies and Glynn, 1968. 


258 VALVIFERA * IDOTEIDAE 


Erichsonella attenuata (Harger, 1873) 
Figure 108B 


DIAGNOSIS 6 11.4 mm, ovigerous 2 12.0 mm. Body dorsally smooth. 
Cephalon lacking middorsal elevation. Antennule reaching only slightly be- 
yond antennal peduncular article 2. Pleotelson with slight marginal bulge in 
anterior half, indicating ventrolateral articulation of uropod. 


RECORDS Connecticut to Miami; Florida, Mississippi, Texas, Gulf of Mex- 
ico; intertidal to 2 m, usually associated with submerged seagrass and algal 


beds. 


REMARKS’ While not recorded in the Florida Keys, this species does reach 
Miami, and continues into the Gulf of Mexico. 


Erichsonella filiformis (Say, 1818) 
Figure 108C 


DIAGNOSIS 6 10.5 mm, ovigerous 2 8.2 mm. Body dorsally with bifid tu- 
bercle on cephalon, and low rounded middorsal tubercle on pereonites. An- 
tennule reaching midlength of antennal peduncular article 3. Basis of per- 
eopods 2—7 with larges tubercles. Pleotelson with distinct lateral shoulder in 


anterior half. 


RECORDS Connecticut to Florida, shallow infratidal to 55 m; Bahamas; 
Turks and Caicos Islands; Puerto Rico; Quintana Roo, Yucatan Peninsula, 
Mexico, 60—109 m; Florida and Texas, Gulf of Mexico. 


Brazil. 


Erichsonella floridana Richardson, 1901 
Figure 108D 


DIAGNOSIS Ovigerous ¢ 10.0 mm. Antennule reaching distal end of anten- 
nal peduncular article 3. Cephalon with strong trifid tubercle. Pereonites 1—7 
each with posteriorly directed spine near posterior margin; pereonites 1—4 
each with lateral spine. Basis of pereopods 2—7 smooth. 


RECORDS Florida Keys, intertidal to 2 m; Florida, Gulf of Mexico, interti- 
dal mud flats. 


Miratidotea 259 


Idotea Fabricius, 1798 


DIAGNOSIS Antennal flagellum multiarticulate. Maxillipedal palp of four or 
five articles. Uropod uniramous. Pleon consisting of two complete and one 
incomplete pleonites plus pleotelson. 


Key to species of Idotea 


ie Postenormargin) of pleotelson, truncate, yo...) je a cuneis.s » (ores eae metallica 
Posterior margin of pleotelson with distinct median lobe ...... balthica 


Idotea balthica (Pallas, 1772) 
Figure 108E 


DIAGNOSIS. 6 24.5 mm, ovigerous 2 13.2—23.5 mm. Anterior margin of 
cephalon concave. Cephalon dorsally smooth. Pereonites evenly convex, 
smooth. Posterior margin of pleotelson with rounded median lobe. 


RECORDS Worldwide in tropical to cold-temperate waters, often on floating 
seaweed, from surface to 357 m. 


Idotea metallica Bosc, 1802 
Figure 108F 


DIAGNOSIS 6 30.0 mm, ovigerous 2? 22.2 mm. Cephalon with sinuous fur- 
row in posterior half. Pereonites 2—4 laterally with rounded convex area close 
to coxae. Posterior margin of pleotelson truncate. 


RECORDS Worldwide in tropical to cold-temperate waters, often on floating 
seaweed, from surface to 200 m. 


Miratidotea Kensley, 1987a 


DIAGNOSIS. Antennal flagellum of single clavate article. Maxillipedal palp 
of four articles. Uropod uniramous. Pleon consisting of two complete and two 
incomplete pleonites plus pleotelson. 


260. VALVIFERA °¢ IDOTEIDAE 


Miratidotea bruscai Kensley, 1987a 
Figure 108G 


DIAGNOSIS Ovigerous 2 13.0 mm. Body parallel sided. Maxillipedal palp 
of four articles, terminal article very short. Pereopods 1—3 increasing in 
length posteriorly, pereopod 4 reduced, shorter than pereopod 5, and with 
dactylus spinelike, pereopods 5—7 increasing in length. Pleotelson consisting 
of two complete and two incomplete pleonites plus pleotelson; latter with 
broadly rounded posterior margin, and with bifid median process situated 
dorsal to posterior oblique-concave area. 


RECORDS Carrie Bow Cay, Belize, 1.5 m, in hollow root-internodes of sea- 
grass Syringodium filiforme. 


Zoogeography 


FAUNAL PROVINCES 


The area under discussion has been divided into several faunal regions or 
provinces, of which the Caribbean, West Indian, and Brazilian are the major 
ones (Briggs, 1974). The extent and boundaries of the provinces have been 
variously defined depending on the group of organisms under discussion. 
Inevitably, zones of overlap exist, but for the purposes of this discussion, the 
following rough limits have been used. 

Brazilian Province: This province stretches from Cape Frio near Rio de 
Janeiro in Brazil to the mouth of the Orinoco River in Venezuela. The out- 
flow of freshwater from the major rivers of this region has probably contrib- 
uted to the isolation of the Brazilian coral reefs and their associated fauna 
from those of the Caribbean. This isolation is demonstrated by the consider- 
able endemism of the Brazilian reef fauna and that of the Caribbean reef 
fauna, with very few species being common to both. 

Caribbean Province: This province has two components, a northern part 
in peninsular Florida, that stretches from around Cape Kennedy on the east 
coast to Tampa or Sanibel Island on the west coast, and a southern compo- 
nent that runs from the mouth of the Orinoco River to around Cabo Rojo or 
Tampico on the gulf coast of Mexico. The northern Gulf of Mexico is ex- 
cluded from this province and is characterized as being warm-temperate, 
rather than subtropical (Briggs, 1974:66). 

West Indian Province: This includes all the islands of the West Indian 
chain, the Bahamas, and the isolated outrider, Bermuda. The West Indian 
Province closely approaches the Caribbean Province in the Yucatan Penin- 
sula to the north, and between Grenada and Trinidad in the south. There is 
also some indication of the isolating effect on the Bahamas of the Florida 
Current through the Straits of Florida. 

It has been suggested, on the basis of the molluscan fauna, that a relict of 
the Neogene Gatunian Province exists around northern Venezuela and Col- 
ombia (Petuch, 1982). While several isopod species have been recorded only 
from this area, these are all described in a single paper that covers a very 
small part of this region (Paul and Menzies, 1971). There is as yet too little 
evidence to explore the idea of this relict fauna further. 


261 


262 ZOOGEOGRAPHY 


ANALYSIS OF THE ISOPOD FAUNA 


In the following discussion, the West Indian and Caribbean provinces are 
treated as one, the isopod faunas offering little evidence to warrant a separate 
treatment of each. 

It is a truism that for any discussion of the zoogeography of an area to have 
meaning, the true extent of the fauna must be known. With the area under 
review, collecting effort has been uneven, and the true faunal composition of 
many regions is still incompletely known. Obviously, any conclusions based 
on such incomplete data are approximate and subject to revision. Neverthe- 
less, certain general patterns or trends emerge when the present isopod fauna 
is broken down into its components. 

The deepwater isopod fauna of the Caribbean (i.e., from deeper than 200 
m) has barely been explored, and little is to be gained from discussing the 
relatively few species known. A list of these deeper dwelling species is in- 
cluded (Table 4). 

Although about 280 shallow-water species are covered by this work, cer- 
tain categories of species must be excluded, for various reasons, before anal- 
ysis can be attempted. Such excluded groups include the species of Oniscidea 
(being essentially terrestrial forms and not part of the marine regime); the 
cymothoid species and the species of Aegidae (being fish parasites for at least 
part of their life history, and whose distribution is complicated by the dis- 
tribution and mobility of the hosts); the limnoriids (being wood-borers whose 
distribution is more a function of the distribution of floating wood); and the 
true cave species (which have a history more reflective of the geological his- 
tory of the area than of the marine regime). The epicarideans have a distribu- 
tion somewhat complicated by the distribution of their crustacean hosts and 
their pelagic epicaridean and cryptoniscan larvae. Nevertheless, the decapod 
hosts of the great majority of species covered here are Caribbean endemics, 
and inclusion of the epicarideans changes very little the overall patterns of 
distribution, as demonstrated by the two figures provided (Figure 109). After 
making these exclusions there remain about 166 species (218 with the epi- 
carideans) that can be broken down into the following components (figures 
in brackets include epicarideans): 

1. True Caribbean/Bahamian species— 124 species, 74.8% [147, 67.5%]. 
These are the species recorded only from the Caribbean and the Bahamas. 
The term endemic is avoided, as too little is known of the actual distribution 
of many species. Of these species, 86 [87] have been recorded from a single 
locality. 

2. Species occurring south of the discussion area, and extending into 
Brazil—5 species, 3.0% [9, 4.1%]. These low numbers indicate that the 


TABLE 4. CARIBBEAN ISOPODS RECORDED FROM DEPTHS GREATER THAN 200 M 


SUBORDER ANTHURIDEA 
Family Paranthuridae 
Neoanthura coeca Menzies, 1956b. South of Jamaica, 1244 m 
SUBORDER ASELLOTA 
Family Dendrotiidae 
Dendrotion hanseni Menzies, 1956b. South of Jamaica, 1244 m 
Family Desmosomatidae 
Desmosoma magnispina Menzies, 1962a. Bay of Panama, 1906 m 
Family Echinothambematidae 
Echinothambema ophiuroides Menzies, 1956a. North of Puerto Rico Trench, 
5104-5122 m 
Family Eurycopidae 
Acanthocope spinosissima Menzies, 1956b. South of Jamaica, 1224 m 
Storthyngura pulchra caribbea (Benedict, 1901). Off Windward Islands, 1256 m 
Storthyngura snanoi Menzies, 1962a. Colombia abyssal plain, 4071 m 
Family Haploniscidae 
Antennuloniscus dimeroceras (Barnard, 1920). North of Puerto Rico Trench, 
5440-5410 m; South Atlantic off South and West Africa, 1400-3921 m; 
off Argentina, 5843 m 
Haploniscus unicornis Menzies, 1956a. North of Puerto Rico Trench, 5104— 
5122 m 
Hydroniscus quadrifrons Menzies, 1962a. North of Puerto Rico Trench, 5271— 
5684 m 
Family Ischnomesidae 
Haplomesus tropicalis Menzies, 1962a. Colombia abyssal plain, 4071 m; off 
South Africa, 2526 m; Mediterranean 
Heteromesus bifurcatus Menzies, 1962a. Colombia abyssal plain, 4071 m 
Ischnomesus armatus Hansen, 1916. North of Puerto Rico Trench, 5494—5477 
m; Davis Straits, 2702 m 
Ischnomesus caribbicus Menzies, 1962a. Off Panama, 1714 m 
Ischnomesus multispinis Menzies, 1962a. Off Panama, 975 m 
Family Janiridae 
Abyssianira dentifrons Menzies, 1956a. North of Puerto Rico Trench, 5104— 
5122 m; off Argentina, 5024-5293 m; off southwest Africa, 4588 m 
Tanirella caribbica Menzies, 1956b. South of Jamaica, 1244 m 
Tanirella vemae Menzies, 1956a. Near Puerto Rico Trench, 5104—5122 m 
Spinianirella serrata Kensley and Heard, 1985. Off Puerto Rico, 350 m 
Family Macrostylidae 
Macrostylis caribbicus Menzies, 1962a. Off Colombia, 2875-2941 m 
Macrostylis minutus Menzies, 1962a. North of Puerto Rico Trench, 5163— 
5494 


(continued ) 


264 ZOOGEOGRAPHY 


TABLE 4. (Continued) 


Macrostylis setifer Menzies, 1962a. North of Puerto Rico Trench, 5477— 
5494 m 
Macrostylis vemae Menzies, 1962a. North of Puerto Rico Trench, 5410— 
5684 m 
Family Mesosignidae 
Mesosignum kohleri Menzies, 1962a. Colombia abyssal plain, 2868-4076 m 
Family Nannoniscidae 
Nannoniscus camayae Menzies, 1962a. Off Panama, 1714 m 
SUBORDER GNATHIIDEA 
Family Gnathiidae 
Akidognathia poteriophora Monod, 1926. Off U. S. Virgin Islands, 914 m. 
SUBORDER VALVIFERA 
Family Arcturidae 
Antarcturus annaoides Menzies, 1956b. South of Jamaica, 1244 m 
Arcturus caribbaeus Richardson, 1901. Off Aves Island, 1360 m 
Arcturus purpureus Beddard, 1886. Off Leeward Islands, 900 m 


Note: Records from deep water around Bermuda are not included. 


great area of mixed-salinity waters resulting from the outflow of the Orinoco, 
Amazon, Tocantins, and Parnaiba rivers form an effective barrier to the 
movement of shallow-water isopod species. 

3. Species having an amphi-Panamic distribution—7 species, 4.2% [8, 
3.7% | (Table 5). In spite of the history of immergence and emergence of the 
Isthmus of Panama, this very small amphi-Panamic component in the Carib- 
bean isopod fauna suggests that most of this fauna has evolved since the last 
emergence of the late Pliocene. Given the limited mobility of most isopod 
species, the Panama Canal seems to have played a minimal role in contribut- 
ing to this component. 

4. Species occurring outside of the western Atlantic (but excluding the 
amphi-Panamic species)—3 species, 1.8% [7, 3.2%]. 

5. The role of the Gulf of Mexico isopod fauna (see Clark and Robertson, 
1982) in the composition of the Caribbean/Bahamian is complex and diffi- 
cult to analyze. One hundred and thirteen species of shallow-water isopods 
have been recorded from the Gulf of Mexico (Table 6). This number would 
indicate that many species remain to be recorded in this region. Of these 113 
species, 61 (54%) have also been reported from the Caribbean region. It is 
therefore possible that there exists a true Gulf of Mexico fauna, whose evolu- 
tion was perhaps spurred by the relative isolation and reduction of the Gulf 


ZOOGEOGRAPHY 265 


Car 74.8% 


GoM/N/Car 7.2% 


GoM/Car 5.4% 


North 3.6% 
Out 1.8% 
Panam 4.2% 


South 3.0% == 


Epicaridea excluded 


Car 67.5%/ 


GoM/N/Car 9.6% 


GoM/Car 6.4% 


South 4.1% 
North 5.5% 
Panam 3.7% Out 3.2% 


Epicaridea included 


Figure 109. Relative proportions of the zoogeographic components of the 
Caribbean isopod fauna, with and without the parasitic Epicaridea. Car, 
Caribbean; Out, extra-western Atlantic; GoM/Car, Gulf of Mexico—Caribbean; 
GoM/N/Car, Gulf of Mexico-Northern-Caribbean; North, northern; Panam, amphi- 
Panamic; South, southern. 


during a low-water stand (100 m below present sea level) during the 
Pleistocene. A significant proportion (about 27 species, 28%) of the Gulf of 
Mexico isopods are known from the eastern coast of the United States north 


266 ZOOGEOGRAPHY 


TABLE 5. SPECIES OF ISOPODS OCCURRING ON BOTH SIDES OF THE ISTHMUS 
OF PANAMA 


*Aega deshaysiana (H. Milne Edwards, _Paradella dianae (Menzies, 1962b) 


1840) Paraleptosphaeroma glynni Buss and 
Anopsilana browni (Van Name, 1936) Iverson, 1981 
Cleantioides planicauda (Benedict, Probopyrus pandalicola (Packard, 
1899) 1879) 
Excirolana braziliensis Richardson, *Rocinela oculata Harger, 1883 
1912 *Rocinela signata Schioedte and 
Excorallana tricornis (Hansen, 1890) Meinert, 1879 
*Nerocila acuminata Schioedte and Uromunna reynoldsi Frankenberg and 
Meinert, 1881 Menzies, 1966 


* fish parasite or fish predator 


of Cape Kennedy, which would indicate a significant cooler-water compo- 
nent. What proportion of originally Gulf species have spread into the Carib- 
bean, and what proportion of Caribbean and temperate east coast species 
have entered the Gulf, cannot yet be assessed, given our incomplete knowl- 
edge of the Gulf fauna. Because of this unresolved situation, three categories 
of species have been separated: species ranging from north of Cape Kennedy 
into the Caribbean—6, 3.6% [12, 5.5%]; species occurring in the Gulf of 
Mexico and the Caribbean—9, 5.4% [14, 6.4%]; species occurring north of 
Cape Kennedy, in the Gulf, and in the Caribbean— 12, 7.2% [21, 9.6%] The 
conclusion that the fauna of the Gulf of Mexico contains an endemic compo- 
nent, a Caribbean component, and a warm-temperate component was also 
reached by Topp and Hoff (1972), in an analysis of the pleuronectiform 
fishes of the Gulf. 


THE BAHAMAS 


The Florida Current flowing through the Straits of Florida has been sug- 
gested as a factor in reducing the movement of shallow-water fauna between 
peninsular Florida and the Florida Keys on the west and the Bahamas on the 
east (Briggs, 1974). Comparison of the number of isopod species on either 
side of the Straits of Florida (13 from the Bahamas, 50 from southern penin- 
sular Florida and the Florida Keys) supports this view. Of the 13 species 
from the Bahamas, only four are “endemic,” three of these being interstitial 
microcerberideans. 


ZOOGEOGRAPHY 267 


TABLE 6. ISOPOD SPECIES OCCURRING IN THE GULF OF MEXICO 


SUBORDER ANTHURIDEA 

*Accalathura crenulata (Richardson, 
1901) 

*Amakusanthura magnifica (Menzies 
and Frankenberg, 1966) 

Cyathura polita (Stimpson, 1855) 

Horoloanthura irpex Menzies and 
Frankenberg, 1966 

Kupellonura formosa (Menzies and 
Frankenberg, 1966) 

* Mesanthura floridensis Menzies and 
Kruczynski, 1983 

* Mesanthura hopkinsi Hooker, 1985 

* Mesanthura pulchra Barnard, 
1925 

Paranthura floridensis Menzies and 
Kruczynski, 1983 

Ptilanthura tricarina Menzies and 
Frankenberg, 1966 

Skuphonura lindae Menzies and 
Kruczynski, 1983 

*Xenanthura brevitelson Barnard, 
1925 

SUBORDER ASELLOTA 

Carpias floridensis Menzies and 
Kruczynski, 1983 

Gnathostenetrioides pugio Hooker, 
1985 

* Joeropsis coralicola Schultz and 
McCloskey, 1967 

* Joeropsis rathbunae Richardson, 
1902 

Mexicope kensleyi Hooker, 1985 

Munnogonium wilsoni Hooker, 1985 

*Pleurocope floridensis Hooker, 
1985 

*Santia milleri (Menzies and Glynn, 
1968) 

*Stenetrium stebbingi Richardson, 
1902 


Uromunna hayesi Robertson, 1978 
*Uromunna reynoldsi Frankenberg 
and Menzies, 1966 


SUBORDER EPICARIDEA 


Allodiplophryxus floridanus Markham, 
1985 

*Aporobopyrina anomala Markham, 
1973 

*Azygopleon schmitti (Pearse, 1932) 

*Bopyrina abbreviata Richardson, 
1904 

*Bopyrione synalphei Bourdon and 
Markham, 1980 

*Cancricepon choprae (Nierstrasz and 
Brender a Brandis, 1925) 

Dactylokepon sulcipes Adkison, 1982 

Eophryxus subcaudalis (Hay, 1917) 

*Gigantione mortenseni Adkison, 
1984b 

Gigantione uberlackerae Adkison, 
1984b 

*Hemiarthrus synalphei (Pearse, 
1950) 

Hyperphrixus castrensis Markham, 
1985 

* Munidion longipedis Markham, 
1975a 

Ovobopyrus alphezemiotes Markham, 
1985 

Parabopyrella mortenseni (Nierstrasz 
and Brender a Brandis, 1929) 

*Parabopyrella richardsonae 
(Nierstrasz and Brender a 
Brandis, 1929) 

Parabopyriscus stellatus Markham, 
1985 

*Probopyria alphei (Richardson, 
1900b) 

Probopyrinella heardi Adkison, 
1984a 


(continued ) 


268 


TABLE 6. (Continued) 


*Probopyrinella latreuticola (Gissler, 
1882) 

Prodajus cf. bigelowiensis Schultz and 
Allen, 1982 

Pseudione cognata Markham, 1985 

Pseudione upogebiae Hay, 1917 

*Schizobopyrina urocaridis (Richardson, 
1904) 

*Stegophryxus hyptius Thompson, 1902 

*Synsynella choprae (Pearse, 1932) 

*Synsynella deformans Hay, 1917 

Synsynella integra Bourdon, 1981 

*Urobopyrus processae Richardson, 1904 


SUBORDER FLABELLIFERA 


*Aega deshaysiana (H. Milne Edwards, 
1840) 

*Aega ecarinata Richardson, 1898 

Aega incisa Schioedte and Meinert, 
1879 

Alcirona krebsii Hansen, 1890 

Ancinus depressus (Say, 1818) 

Anilocra acuta Richardson, 1910 

Anilocra laticauda H. Milne Edwards, 
1840 

*Bathynomus giganteus A. Milne 
Edwards, 1879 

* Cassidinidea ovalis (Say, 1818) 

Ceratothoa transversa (Richardson, 
1900b) 

Cirolana borealis Lilljeborg, 1851 

*Cirolana obtruncata Richardson, 1901 

*Cirolana parva Hansen, 1890 

Conilera cylindracea (Montagu, 1804) 

*Cymothoa caraibica Bovallius, 1885 

*Cymothoa excisa Perty, 1833 

*Cymothoa oestrum (Linnaeus, 1793) 

*Cerceis carinata Glynn, 1970 

*Furydice convexa Richardson, 1900b 


Eurydice littoralis (Moore, 1901) 
*Eurydice piperata Menzies and 
Frankenberg, 1966 


*Excirolana braziliensis Richardson, 
1912a 

*Excirolana mayana (Ives, 1891) 

*Excorallana antillensis (Hansen, 
1890) 

Excorallana mexicana Richardson, 
1905a 

*Excorallana tricornis (Hansen, 
1890) 

*Harrieta faxoni (Richardson, 
1905a) 

*Limnoria tuberculata Sowinsky, 
1884 

Lironeca ovalis (Say, 1818) 

*Lironeca redmanni Leach, 1818 

Lironeca texana Pearse, 1952 

Lironeca tropicalis Menzies and 
Kruczynski, 1983 

*Nalicora rapax Moore, 1901 

*Nerocila acuminata Schioedte and 
Meinert, 1881 

Olencira praegustator (Latrobe, 1802) 

*Paracerceis caudata (Say, 1818) 

*Paradella dianae (Menzies, 1962b) 

Paradynamene benjamensis 
Richardson, 1905 

*Politolana polita (Stimpson, 1853) 

*Rocinela insularis Schioedte and 
Meinert, 1879 

*Rocinela oculata Harger, 1883 

*Rocinela signata Schioedte and 
Meinert, 1879 

*Serolis mgrayi Menzies and 
Frankenberg, 1966 

*Sphaeroma quadridentata Say, 1818 

*Sphaeroma terebrans Bate, 1866 


SUBORDER GNATHIIDEA 


Gnathia floridensis Menzies and 
Kruczynski, 1983 
SUBORDER MICROCERBERIDEA 
Microcerberus mexicanus Pennak, 1958 


SUBORDER VALVIFERA 

Antarcturus floridanus (Richardson, 
1900b) 

Arcturella bispinata Menzies and 
Kruczynski, 1983 

Arcturella spinata Menzies and 
Kruczynski, 1983 

Astacilla lauffi Menzies and 
Frankenberg, 1966 

Chiridotea excavata Harper, 


ZOOGEOGRAPHY 269 


Edotea lyonsi (Menzies and 
Kruczynski, 1983) 

Edotea montosa (Stimpson, 1853) 

Edwinjoycea horologium Menzies and 
Kruczynski, 1983 

*Erichsonella attenuata (Harger, 
1873) 

*Frichsonella filiformis (Say, 1818) 

*Erichsonella floridana Benedict, 
1901 


1974 Erichsonella isabelensis Menzies, 
*Cleantioides planicauda (Benedict, 1951b 
1899) *Idotea metallica Bosc, 1802 


* species also occurring in the Caribbean 
Note: Records for the Gulf of Mexico have been assembled from published 
literature; in most cases, actual material has not been examined. 


BERMUDA 


Twenty-nine species of isopods have been recorded from Bermuda (Table 7). 
Of these, nine are endemics (three being cave forms). The remaining 20 
species have all been recorded from the Caribbean region, indicating a strong 
subtropical connection, in spite of the relatively high latitude (32°15'N). Al- 
though Bermuda is of Eocene or Oligocene age, the tropical fauna was prob- 
ably decimated by the low temperatures of the last Pleistocene glaciation 
(Briggs, 1974:76). 


CAVE ISOPODS 


With the expanding efforts of cave divers, more and more true stygobiont 
forms are being found. Concurrently, discussion of the origin of cave fauna 
has spurred several theories, all invoking the geological history of the Carib- 
bean area. 

Among the isopods, cave forms have been found in four suborders, the 
Asellota, Anthuridea, Flabellifera, and Microcerberidea. Two valuable dis- 
cussions on the origin of cave crustaceans may be found in Stock (1986) and 
Wagele (1985). 

The only true cave asellote, Atlantasellus cavernicolus Sket, was collected 


from Bermuda. 


270 ZOOGEOGRAPHY 


Alcirona krebsi Hansen, 1890 

*Anthomuda stenotelson Schultz, 1979 

*Apanthura harringtoniensis Wagele, 
1981 

*Arubolana aruboides (Bowman and 
Iliffe, 1983) 

*Atlantasellus cavernicolus Sket, 1979 

Bopyrissa wolffi Markham, 1978 

Cancricepon choprae (Nierstrasz and 
Brender a Brandis, 1925) 

Carpias bermudensis Richardson, 1902 

*Carpias minutus (Richardson, 1902) 

*Colanthura tenuis Richardson, 1902 

Colopisthus parvus Richardson, 1902 

*Curassanthura bermudensis Wagele, 
1985 

Dynamenella perforata (Moore, 1901) 

Eurydice personata Kensley, 1987b 


TABLE 7. ISOPOD SPECIES OCCURRING AT BERMUDA 


Excorallana quadricornis (Hansen, 1890) 

Joeropsis rathbunae Richardson, 1902 

Leidya bimini Pearse, 1951 

Paracerceis caudata (Say, 1818) 

Paranthura infundibulata Richardson, 
1902 

Pendanthura tanaiformis Menzies and 
Glynn, 1968 

Parathelges piriformis Markham, 1972b 

Parathelges tumidipes Markham, 1972b 

Probopyrinella latreuticola (Gissler, 
1882) 

Pseudione affinis (Sars, 1882) 

*Stegias clibanarii Richardson, 1904 

Stenetrium stebbingi Richardson, 1902 

Stenobermuda acutirostrata Schultz, 1979 

Synsynella choprae (Pearse, 1932) 

Synsynella deformans Hay, 1917 


* recorded only from Bermuda 


The anthuridean cave representatives are found in two families: the genus 
Curassanthura Kensley in the Paranthuridae, and the genus Cyathura subgenus 
Stygocyathura Botosaneanu and Stock in the Anthuridae (see Figure 110). 

Three species of Curassanthura are known, one each from Curagao, Ber- 
muda, and Lanzarote in the Canary Islands. Curassanthura halma Kensley, 
from Curacao, is an interstitial form found in hypersaline waters. Curas- 
santhura bermudensis Wagele was found in water of about 26%o salinity. The 
Lanzarote species, C. canariensis Wagele, came from seawater in a lava cave. 
Wagele (1985) suggests that this amphi-Atlantic distribution of Curassanthura— 
is the result of plate tectonics separating an ancestral hypogean progenitor 
that had a Tethyan distribution. 

The genus Cyathura has representatives in the sea, in estuarine-brackish 
habitats, and in freshwater caves, and is found in the Atlantic, Indian, and 
Pacific oceans. This widespread distribution suggests a very long history for 
the genus. Using the morphology of the male copulatory stylet, Wagele 
(1985) suggests that marine ancestors, having a Tethyan distribution, en- 
tered freshwater interstitial habitats. The series of regressions of sea level 
during the Pleistocene probably served further to isolate these freshwater 
forms. 


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ZOOGEOGRAPHY DAs 


The flabelliferan family Cirolanidae contains five stygobiont genera in the 
Caribbean: Anopsilana, Arubolana, Bahalana, Creaseriella, and Haptolana (Figure 
111). Six other genera are known from the North American continent: 
Antrolana, Cirolanides, Mexilana, Speocirolana, Sphaerolana, and Troglocirolana, all 
of which, except Antrolana from the Appalachian Valley of Virginia, occur in 
Mexico and Texas (see Notenboom, 1981). A few of these forms occur in 
brackish water, but most are found in freshwater of caves. Cave cirolanids 
are also known from Palau, North and East Africa, Madagascar, Bulgaria, 
Greece, Jugoslavia, Israel, France, and Spain. This widespread distribution 
again suggests a Tethyan marine origin, with dispersal and isolation due to 
sea regressions. 

The suborder Microcerberidea and the asellotan family Microparasellidae 
contain almost entirely interstitial forms, although few occur in caves. At 
least two genera, Microcerberus and Angliera, have very widespread distribu- 
tions and are known from marine, brackish-water, and freshwater habitats, 
and may well have a history similar to that of Cyathura. 


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Appendix 


Since the manuscript of this work was completed and sent to press, a few 
papers have appeared either describing new species, mentioning new 
records for the Caribbean and associated areas, or instituting a major 
new taxon. It was thought useful to include these, if only in an appendix, 
to make the work as current as possible. The relevant taxa are listed 
alphabetically, with the full citation given below. 


Antheluridae Poore and Lew Ton, 1988 
Poore, G. C. B., and H. M. Lew Ton. 1988. Antheluridae, a new 
family of Crustacea (Isopoda: Anthuridea) with new species from 
Australia. Journal of Natural History 22:489—506. 
Within the geographical area covered by this work, only Anthomuda 
belongs to this new family. 


Aporobopyrus collardi Adkison, 1988 
Adkison, D. L. 1988. Pseudione parviramus and Aporobopyrus collard, 
two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf 
of Mexico. Proceedings of the Biological Society of Washington 
101(3):576—584. 


Booralana tricarinata Camp and Heard, 1988 

Camp, D. K., and R. W. Heard. 1988. Booralana tricarinata, a new 
species of isopod from the western Atlantic Ocean (Crustacea: Iso- 
poda: Cirolanidae). Proceedings of the Biological Society of Washington 
101 (3):603-613. 

Originally recorded from the outer shelf and upper slope off the 
Little Bahama Bank and the Antilles Islands in 110—610 m, this 
species has since been recorded off Haiti in 620 m. 


Bythognathia yucatanensis Camp, 1988 


Camp, D. K. 1988. Bythognathia yucatanensis, new genus, new species, 
from abyssal depths in the Caribbean Sea, with a list of gnathiid 


275 


276 APPENDIX 


species described since 1926 (Isopoda: Gnathiidae). Journal of Crust- 
acean Biology 8(4):668—678. 


Edotea samariensis Miller, 1988 
Miller, H. G. 1988. Idoteidae aus N-Kolumbien mit Beschreibung 
von Edotea samariensis n. sp. (Crustacea: Isopoda: Valvifera). 
Senckenbergiana biologica 68(4/6):407—4 12. 


Gnathia johanna Monod, 1926 
Miller, H. G. 1988. Redescription of Gnathia johanna, 1926 (Iso- 
poda) from St. John, Virgin Islands. Bulletin Zoologisch Museum, Uni- 
versiteit van Amsterdam 11(15):129—133. 


Phycolimnoria bacescui Ortiz and Lalana, 1988 
Ortiz, M., and R. Lalana. 1988. Una nueva especie del genero Phy- 
colimnoria (Isopoda, Limnoriidae) de aguas cubanas. Revista de Inves- 
tigaciones Marinas, La Habana 9(2):37—42. 


Pseudione parviramus Adkison, 1988 
Adkison, D. L. 1988. Pseudione parviramus and Aporobopyrus collardi, 
two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf 
of Mexico. Proceedings of the Biological Society of Washington 
101(3):576—584. 


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292 LITERATURE CITED 


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Index 


abbreviata, Bopyrina, 110, 267 
Abudefduf saxatilis, 171, 175, 186 
abudefdufi, Anilocra, 171, 175 
Abyssianira dentifrons, 263 
Acanthocope spinosissima, 263 
acanthophora, Domecia, 111 
Acanthopleura granulata, 215 
acanthura, Anopsilana, 124, 125 
acanthuri, Anilocra, 171, 175, 176 
Acanthurus 

bahianus, 171, 176 

chirurgus, 171, 176 
acanthurus, Macrobrachium, 112 
Accalathura,64 

crenulata, 64, 65, 267 

setosa, 64, 65 
Achelion occidentalis, 110 
acuminata, Nerocila, 171, 172, 173, 190, 266, 

268 

forma acuminata, 190 

forma aster, 190 
acuta, Anilocra, 268 
acutirostrata, Stenobermuda, 106, 270 
acutirostris, Processa, 113 
acutitelson, Dynamenella, 213, 214 
Aega, 116, 117 

antillensis, 117 

incisa, 268 
Aega (Aega), 116, 117 

deshaysiana, 117, 266, 268 

ecarinata, 117, 268 
Aega (Rhamphion), 116, 117 

dentata, 117, 119 

tenuipes, 117, 119 
Aegidae, 115, 116, 262 
Aetobatus narinari, 166 
affinis 

Pseudione, 113, 270 

Upogebia, 112 
Agaricia, 88, 166 
agaricicola, Metacirolana, 153, 154 
Agarna, 170, 173 

cumulus, 173 


Agelas, 88 
Akidognathia poteriophora, 264 
alba, Exosphaeroma, 229 
albidoida, Cirolana, 132, 133 
Alcirona, 157, 158 

insularis, 158 

krebsii, 158, 268, 270 
algicola, Carpias, 82, 83, 87 
Allodardanus bredini, 112 
Allodiplophryxus floridanus, 267 
alphei, Probopyria, 112, 267 
Alpheus 

armillatus, 112 

formosus, 111, 112 

heterochaelis, 112 

normanni, 112 

viridari, 112 
alphezemiotes, Ovobopyrus, 267 
Alutera schoepfi, 171, 190 
Amakusanthura, 17, 18 

geminsula, 18 

lathridia, 18, 20 

magnifica, 18, 20, 267 

signata, 18, 21 

significa, 18, 23 
amazonicum, Macrobrachium, 112 
Ambidexter symmetricus, 113 
americana, Dasyatis, 122, 166 
americanus, Periclimenes, 111, 113 
Amphiroa, 219 
amyle, Licranthura, 43 
analis, Lutjanus, 122, 172, 183 
Anchoa lamprotaenia, 171, 187 
Ancininae, 204 
Ancinus, 205 

belizensis, 205 

brasiliensis, 205, 206 

depressus, 268 
andrewsi, Paralimnoria, 199 
Angliera, 90, 91, 273 

dubitans, 91 

psamathus, 91 

racovitzai, 91 


293 


294 INDEX 


angulata 

Dynamene, 214 

Dynamenella, 214 
angustifrons, Hexapanopeus, 11] 
Anilocra, 170, 174, 175 

abudefdufi, 171, 175 

acanthuri, 171, 175, 176 

acuta, 268 

chaetodontis, 171, 175, 177 

chromis, 172, 175, 177 

haemuli, 172, 173, 175, 177 

holacanthi, 172, 175, 179 

holocentri, 175, 179 

laticauda, 268 

myripristis, 172, 175, 180 

partiti, 173, 175, 180 
annandalei, Sphaeroma, 234 
annaoides, Antarcturus, 264 
annulicornis, Pandalus, 113 
annulipes, Pagurus, 113 
anomala, Aporobopyrina, 110, 267 
anops, Creaseriella, 137 
Anopsilana, 124, 273 

acanthura, 124, 125 

browni, 124, 125, 266 

crenata, 124, 125 

cubensis, 124, 126 

Jonesi, 124, 127 

radicicola, 124, 127 
Antarcturus 

annaoides, 264 

Sloridanus, 269 
Antennuloniscus dimeroceras, 263 
Antheluridae, 275 
Anthomuda, 17, 23, 291 

stenotelson, 23, 270 
Anthuridae, 16, 270 
Anthuridea, 2, 14, 15, 16, 244, 263, 269 
antiguai, Plesionika, 113 
antillense, Exosphaeroma, 229, 230 
antillensis 

Aega, 117 

Dromidia, 111 

Excorallana, 161, 162, 268 

Paranthura, 69 
Antrolana, 273 
Apanthura, 17, 25 

cracenta, 25 

crucis, 25, 26 

harringtoniensis, 25, 26, 270 
Apanthuroides, 17, 26 

millae, 27 
Aphysina fistularis, 223 


Apogon 

lachneri, 171, 188 

maculatus, 171, 193 

townsendi, 171, 193 
Apogonidae, 187 
Aporobopyrina, 110 

anomala, 110, 267 
Aporobopyrus, 110 

collardi, 275 

curtatus, 110 
arbuscula, Oculina, 88 
Archosargus probatocephalus, 122 
Arcturella, 252 

bispinata, 252, 269 

Spinata, 252, 269 
Arcturidae, 251, 252, 264 
Arcturus 

caribbaeus, 264 

purpureus, 264 
arenatus, Priacanthus, 173, 183 
Argeia, 110 

atlantica, 110 
Argeiinae, 107 
Arius felis, 171, 190 
Armadilloniscus, 247 

ninae, 247 
armatus 

Ischnomesus, 263 

Petrolisthes, 110 
armillatus, Alpheus, 112 
arndti, Dies, 207 


aruboides, Arubolana, 144, 145, 270 


Arubolana, 144, 273 

aruboides, 144, 145, 270 

imula, 144, 145 

Parvioculata, 144, 145 
ascensionis, Holocentrus, 172, 179 
Aselloidea, 75 


Asellota, 15, 15, 73, 263, 267, 269 


Astacilla, 252 
cymodocea, 252, 253 
lasallae, 252, 253 
lauffi, 269 
regina, 252, 253 

Astalione, 110 
cruciaria, 110 


aster, forma, Nerocila acuminata, 190 


Astrapogon stellatus, 171, 188 
Asymmetrione, 110 

clibanarii, 110 

desultor, 110 
Athelginae, 107 
Atherinidae, 187 


Atlantasellidae, 75 
Atlantasellus, 75 

cavernicolus, 75, 269, 270 
atlantica 

Argeia, 110 

Megalops, 172, 183 
attenuata, Erichsonella, 256, 258, 269 
aurolineatum, Haemulon, 172, 178 
Azygopleon, 110 

schmitti, 110, 267 
bacescui, Phycolimnoria, 276 
Bagatus, 83 
Bahalana, 124, 127, 128, 273 

cardiopus, 128 

geracei, 128 

mayana, 128 
bahianus, Acanthurus, 171, 176 
bajonado, Calamus, 122 
Balanopleon, 110 

tortuganus, 110 
Balistes vetula, 122 
balthica, Idotea, 259 
barbadensis, Micropanope, 111 
barbarae, Geogerceis, 215 
barnardi 

Dies, 207 

Ischyromene, 218 

Paranthura, 69, 71 
barracuda, Sphyraena, 122 
barrerae, Cymodoce, 227 
Bathynomus, 123, 129 

giganteus, 131, 268 
Batrachoides surinamensis, 171, 190 
baudiniana, Ligia, 247, 249 
beethoveni, Gnathia, 238, 239 
belizensis, Ancinus, 205 
Belonidae, 187 
benjamensis, Paradynamene, 268 
berbicensis, Excorallana, 161, 162 
bermudensis 

Carpias, 82, 83, 270 

Curassanthura, 67, 270 

Hemiramphus, 188 
bicornutus, Microphrys, 110 
bifasciatus, Joeropsis, 88 
bifurcatus, Heteromesus, 263 
bigelowiensis, cf., Prodajus, 268 
bimini, Leidya, 111, 270 
bispinata, Arcturella, 252, 269 
bivittata, Mesanthura, 47 
blackfordi, Lutjanus, 122 
bohlkeorum, Mothocya, 171, 173, 187, 188 
bonaci, Mycteroperca, 122 


INDEX 


bonairensis, Pagurus, 110, 113 
bonelli, Macrobrachium, 112 
bonnieri, Pandalus, 113 
Booralana tricarinata, 275 
Bopyrella, 110 

harmopleon, 110 
Bopyridae, 107, 114 
Bopyrina, 110 

abbreviata, 110, 267 
Bopyrinae, 107 
Bopyrinella, 110 

thorii, 110 
Bopyrione, 110 

synalphei, 110, 267 
Bopyrissa, 110 

wolffi, 110, 270 
Bopyroidea, 107 
Bopyrophryxinae, 107 
borealis, Cirolana, 268 
Bothus lunatus, 122 
bousfieldi, Synalpheus, 110 
bowmani 

Renocila, 173, 191 

Stenetrium, 100 
brachydactylus, Carpias, 82, 84 
brasiliensis 

Ancinus, 205, 206 

Hemiramphus, 172, 184 
braziliensis, Excirolana, 150, 266, 268 
bredini, Allodardanus, 112 
brevicarpus, Synalpheus, 110 
brevidactylus, Pagurus, 112, 113 
brevipes, var., Paracerceis caudata, 219 
brevitelson, Xenanthura, 62, 267 
brooksi, Synalpheus, 110, 111, 113 
browni, Anopsilana, 124, 125, 266 
bruscai, Miratidotea, 260 
buccanella, Lutjanus, 122 
Bythognathia yucatanensis, 275 
Cabirops, 110 
Calamus 

bajonado, 122 

calamus, 122 

penna, 122 
calamus, Calamus, 122 
Callispongia plicifera, 221 
Calyptolana, 124, 131 

hancocki,132 
camayae, Nannoniscus, 264 
canaliculata, Processa, 113 
canariensis, Curassanthura, 270, 
Cancricepon, 110 

choprae, 110, 267, 270 


295 


296 INDEX 


Cancrion carolinus, 111 
capistratus, Chaetodon, 171, 177 
caraibica, Cymothoa, 182, 268 
Caranx, 122, 171 

hippos, 171, 183 

latus, 171, 183 

ruber, 171, 183 
carbonarium, Haemulon, 172, 178 
carcinus, Macrobrachium, 112 
cardiopus, Bahalana, 128 
caribaeus, Dactylokepon, 111 
caribbaeus, Arcturus, 264 
caribbea, Storthyngura pulchra, 263 
caribbica 

Tanirella, 263 

Malacanthura, 45 
caribbicus 

Ischnomesus, 263 

Macrostylis, 263 
caribea, Uromunna, 94 
carinata, ‘Cerceis,‘ 211, 268 
carolii, Metaphrixus, 111 
carolinense, Tozeuma, 112 
carolinus, Cancrion, 111 
Carpias, 82, 83 

algicola, 82, 83, 87 

bermudensis, 82, 83, 270 

brachydactylus, 82, 84 

floridensis, 267 

harrietae, 82, 84 

minutus, 82, 84, 270 

punctatus, 82, 85 

serricaudus, 82, 87 

triton, 82, 87 
Cassidinidea, 207, 208 

mosaica, 208 

ovalis, 207, 208, 268 
Cassidininae, 204, 207 
castrensis, Hyperphrixus, 267 
caudata, Paracerceis, 219, 268, 270 
Caulerpa, 219 
cavalla, Scomberomorus, 173, 187 
cavernicolus, Atlantasellus, 75, 269, 270 
cephalus, Mugil, 172, 190 
Ceratothoa, 170, 180 

deplanata, 180 

transversa, 268 
Cerceis, 210, 211 

carinata, 211, 268 
Chaetodipterus faber, 171, 190 
Chaetodon 

capistratus, 171, 177 

ocellatus, 171, 177 


sedentarius, 171, 177 
striatus, 171, 177 


chaetodontis, Anilocra, 171, 175, 177 


Chalixanthura, 17, 27 

lewisi, 27, 29 

scopulosa, 27, 29 
Chilomycterus schoepfi, 172, 190 
Chiridotea excavata, 269 


chirurgus, Acanthurus, 171, 176 


Chiton 

marmoratus, 229 

tuberculatus, 229 
choprae 

Cancricepon, 110, 267, 270 

Synsynella, 113, 268, 270 
Chromis 

cyaneus, 172, 177 

multilineatus, 172, 177 
chromis, Anilocra, 172, 175, 177 
chrysargyreum, Haemulon, 172, 178 
chrysoptera, Orthopristis, 172, 183 
chrysurus, Ocyurus, 172, 183 
ciliatus, Monacanthus, 172, 190 
circumsaltanus, Loki, 111 
Cirolana, 123, 132 

albidoida, 132, 133 

borealis, 268 

crenulitelson, 132, 133 

minuta, 132, 135 

obtruncata, 132, 135, 268 

parva, 132, 135, 268 
Cirolanidae, 1, 115, 122, 123, 139, 273 
Cirolanides, 273 
Cirolaninae, 123, 139 
cirratum, Ginglymostoma, 122, 158 
clarkae, Phycolimnoria, 201 
Clastotoechus vanderhorsti, 110 
Cleantioides, 255, 256 

occidentalis, 256 

planicauda, 256, 266, 269 
clibanarii 

Asymmetrione, 110 

Stegias, 113, 270 
Clibanarius 

tricolor, 110, 112, 113 

vittatus, 110, 112 
coeca, Neoanthura, 263 
cognata, Pseudione, 268 
cohenae, Paracerceis, 219, 220 
Colanthura, 64, 65 

tenuis, 65, 270 
colini, Renocila, 171, 191 
collardi, Aporobopyrus, 275 


Colopisthus, 144, 146 

parvus, 147, 270 
confixa, Cortezura, 31 
Conilera cylindracea, 268 
Conilerinae, 123, 139 
conklini, Phaeoptyx, 173, 188 
constricta, Munida, 112 
convexa, Eurydice, 147, 148, 149, 268 
coralicola, Joeropsis, 88, 267 
Corallanidae, 115, 157 
corallicola, Minyanthura, 53 
corallinus, Pylopagurus, 112 
Cortezura, 17, 29 

confixa, 31 

penascoensis, 31 
cracenta, Apanthura, 25 
crassa, Virganthura, 73 
Creaseriella, 124, 137, 273 

anops, 137 
crenata, Anopsilana, 124, 125 
crenulata, Accalathura, 64, 65, 267 
crenulitelson, Cirolana, 132, 133 
cromis, Pogonias, 173, 190 
cruciaria, Astalione, 110 
crucis, Apanthura, 25, 26 
cruentatus, Epinephelus, 172, 179 
crumenophthalmus, Selar, 173, 183 
Cryptoniscoidea, 107 
Crytoniscidae, 107, 109 
cubana, Cyathura (Cyathura), 33 
cubense, Munidion, 111 
cubensis 

Anopsilana, 124, 126 

Rocinela, 119, 120 
cuborientalis, Cyathura (Stygocyathura), 34, 35 
culebrae, Vandeloscia, 247, 251 
cumanensis, Malacanthura, 45 
cumulus, Agarna, 173 
cuprea, Diopatra, 256 
curacaoensis, Hippolyte, 110 
Curassanthura, 64, 67, 270 

bermudensis, 67, 270 

canariensis, 270 

halma, 67, 68, 270 
curassavica, Cyathura (Stygocyathura), 35 
curri, Haliophasma, 41 
curtatus, Aporobopyrus, 110 
cuvieri, Galeocerdo, 122 
cyaneus, Chromis, 172, 177 
Cyathura, 17, 31, 270, 273 

polita, 267 

(Cyathura), 31 

cubana, 33 


INDEX 


(Stygocyathura), 31, 33, 35, 270 
cuborientalis, 34, 35 
curassavica, 35 
hummelincki, 35 
motasi, 35, 36 
orghidani, 35, 36 
parapotamica, 35, 36 
salpiscinalis, 35, 38 
sbordonii, 35, 38 
specus, 35, 38 
univam, 35, 38 

Cyclograpsus interger, 111 
cylindracea, Conilera, 268 
Cymodoce, 226, 227 

barrerae, 227 

ruetzleri, 227 

Cymodocea, 223, 253 
cymododea, Astacilla, 252, 253 
Cymothoa, 170, 172, 182 
caraibica, 182, 268 
excisa, 172, 173, 182, 268 


oestrum, 171, 172, 173, 182, 183, 268 


Cymothoidae, 115, 169, 170 
Cynoscion, 172, 183 
nebulosus, 172, 183 
Dactylokepon 
caribaeus, 111 
sulcipes, 267 
Dajidae, 107 
Dardanus fucosus, 1122 
Dasyatis americana, 122, 166 
decorata, Mesanthura, 53 


deformans, Synsynella, 110, 113, 268, 270 


delaneyi, Excorallana, 160, 161 
Dendrotiidae, 263 
Dendrotion hanseni, 263 
dentata, Aega, 117, 119 
dentifrons, Abyssianira, 263 
deplanata, Ceratothoa, 180 
depressa, Panoplax, 111 
depressus, Ancinus, 268 
deshaysiana, Aega, 117, 266, 268 
Desmosoma magnispina, 263 
Desmosomatidae, 263 
destructor, Sphaeroma, 235 
desultor, Asymmetrione, 110 
dianae, Paradella, 224, 266, 268 
Dicropleon 

periclimenis, 111 
Dictyota, 219 
Dies, 207 

arndti, 207 

barnardi, 207 


297 


298 INDEX 


dimeroceras, Antennuloniscus, 263 floridana, 256, 258, 269 
diminuta, Exosphaeroma, 229, 231 isabelensis, 269 
diogenes, Petrochirus, 110 Eriphia gonagra, 111 
Diopatra cuprea, 256 Eubranchiatae, 203 
Diplophryxus, 111 Eurycopidae, 263 
Discerceis, 210, 211 Eurydice, 143, 147 
linguicauda, 213 convexa, 147, 148, 149, 268 
dispar, Paraliomera, 111 littoralis, 148, 149, 268 
distorta, Leidya, 111 personata, 147, 149, 270 
Domecia piperata, 147, 149, 268 
acanthophora, 111 Eurydicinae, 123, 139, 143 
hispida, 111 excavata, Chiridotea, 269 
Dromidia antillensis, 111 Excirolana, 144, 149, 150 
dubitans, Angliera, 91 braziliensis, 150, 266, 268 
Dynamene angulata, 214 mayana, 150, 153, 268 
Dynamenella, 210, 213, 214, 224 excisa, Cymothoa, 172, 173, 182, 268 
acutitelson, 213, 214 Excorallana, 157, 159, 161 
var. glabrothorax, 214 antillensis, 161, 162, 268 
var. typica, 214 berbicensis, 161, 162 
angulata, 214 delaneyi, 160, 161 
perforata, 213, 215, 270 fissicauda, 161, 162 
quadrilirata, 213, 215 mexicana, 160, 161, 268 
Dynameninae, 204, 210, 211 oculata, 161, 163 
ecarinata, Aega, 117, 268 quadricornis, 161, 165, 270 
Echinothambema ophiuroides, 263 sexticornis, 161, 165 
Echinothambematidae, 263 subtilis, 160 
edithae, Paracerceis, 218, 221 tricornis, 266, 268 
Edotea occidentalis, 167 
lyonsi, 269 tricornis, 161, 165 
montosa, 269 warmingti, 161, 167 
samariensis, 276 exilipes, Palaemonetes, 113 
edulis, Processa, 113 Exocoettus, 172, 184 
edwardsi, Plesionika, 113 Exosphaeroma, 226, 229, 231 
Edwinjoycea horologium, 252, 269 alba, 229 
eglanteria, Raja, 122 antillense, 229, 230 
Eisothistos, 16, 38, 39 diminuta, 229, 231 
petrensis, 39 productatelson, 229, 231 
teri, 39 yucatanum, 229, 231 
ensis, Plesionika, 113 exotica, Ligia, 247, 249 
Entoniscidae, 107, 109 Jaber, Chaetodipterus, 171, 190 
Entophilinae, 107 fasciata, Mesanthura, 47, 49 
Eophrixus subcaudalis, 111, 267 faustinum, Macrobrachium, 112 
Epicaridea, 4, 14, 107, 267 Jaxoni, Harrieta, 232, 268 
Epinephelus, 122, 172, 183 felis, Arius, 171, 190 
cruentatus, 172, 179 filiforme, Syringodium, 253, 260 
Julvus, 172, 179 filiformis, Erichsonella, 256, 258, 269 
guttatus, 172, 179 fimbriata 
itajara, 122, 172, 190 Pleurocryptella, 112 
morio, 122 Processa, 113 
Erichsonella, 255, 256, 257 Jissicauda, Excorallana, 161, 162 
attenuata, 256, 258, 269 fistularis, Aphysina, 223 
filiformis, 256, 258, 269 Flabellifera, 2, 14, 114, 115, 268, 269 


tropicalis, 257 flavolineatum, Haemulon, 122, 172, 178 


flinti, Munida, 111 
floridana 

Erichsonella, 256, 258, 269 

Pleurocrypta, 112 
floridanus 

Allodiplophryxus, 267 

Antarcturus, 269 

Thor, 110, 111 
Sloridensis 

Carpias, 267 

Gnathia, 268 

Mesanthura, 53, 267 

Paranthura, 69, 71, 267 

Pleurocope, 98, 267 
foetens, Synodus, 173, 183 
foliatus, Parathelges, 112 
formosa, Kupellonura, 267 
formosus, Alpheus, 111, 112 
fritzmuelleri, Synalpheus, 110, 111 
fucorum, Latreutes, 112 
fucosus, Dardanus, 112 
fulvus, Epinephelus, 172, 179 
furcifer, Paranthias, 173, 179 
Galathea rostrata, 112 
galathinus, Petrolisthes, 110 
Galeocerdo cuvieri, 122 
geminsula, Amakusanthura, 18 
Geocerceis, 210, 215 

barbarae, 215 
geracei, Bahalana, 128 
Gerres rhombeus, 172, 187 
giardi, Synalpheion, 113 
giganteus, Bathynomus, 131, 268 
Gigantione 

mortenseni, 111, 267 

uberlackerae, 267 
Ginglymostoma cirratum, 122, 158 


glabrothorax, var., Dynamenella acutitelson, 214 


Glossobius, 170, 172, 183, 184 
hemiramphi, 172, 184 
impressus, 172, 184 

glynni 
Paracerceis, 218, 221 
Paraleptosphaeroma, 210, 266 

Gnathia, 238 
beethoveni, 238, 239 
floridensis, 268 
gonzalezi, 238, 239 
johanna, 238, 239, 276 
magdalenensis, 238, 239 
puertoricensis, 238, 239 
rathi, 238, 241 
samariensis, 238, 241 


INDEX 


triospathiona, 238, 241 

velosa, 238, 243 

virginalis, 238, 243 
Gnathiidae, 237, 238 
Gnathiidea, 14, 236, 238, 268 
Gnathostenetroides, 77 

laodicense, 78 

pugio, 77, 267 
Gnathostenetroididae, 77 
Gnathostenetroidoidea, 77 
gonagra, Eriphia, 111 
gonzalezi, Gnathia, 238, 239 
goodei, Synalpheus, 110, 111 
gracilis, Natatolana, 140 
granulata, Acanthopleura, 215 
guttatus, Epinephelus, 172, 179 
haemuli, Anilocra, 172, 173, 175, 177 
Haemulon 

aurolineatum, 172, 178 

carbonarium, 172, 178 

chrysargyreum, 172, 178 

flavolineatum, 122, 172, 178 

macrostomum, 172, 179 

plumiert, 172, 179 

sciurus, 172, 179 

steindachneri, 122 
halia, Metacirolana, 153, 154 
Halimeda, 88, 90, 219, 223 
Haliophasma, 17, 41 

curri, 41 

irmae, 43 

valeriae, 41, 42 
halma, Curassanthura, 67, 68, 270 
Halodule, 232 
hancocki, Calyptolana, 132 
hanseni, Dendrotion, 263 
Haplomesus tropicalis, 263 
Haploniscidae, 263 
Haploniscus unicornis, 263 
Haptolana, 124, 137, 273 

somala, 138 

trichostoma, 138 
harmopleon, Bopyrella, 110 
Harrieta, 226, 232 

faxoni, 232, 268 
harrietae, Carpias, 82, 84 
harringtoniensis, Apanthura, 25, 26, 270 
harrisii, Rithropanopeus, 111 
hayesi, Uromunna, 267 
heardi, Probopyrinella, 267 
Hemiarthrinae, 107 
Hemiarthrus synalphei, 111, 267 
Hemibranchiatae, 203 


209 


300 INDEX 


hemiramphi, Glossobius, 172, 184 
Hemiramphidae, 187 
Hemiramphus 

bermudensis, 188 

brasiliensis, 172, 184 
hemphilli, Synalpheus, 110, 111 
hendleri, Pendanthura, 56 
herbstii, Panopeus, 111 
herrerai, Microcharon, 93 
heterocarpus, Plesionika, 113 
heterochaelis, Alpheus, 112 
Heteromesus bifurcatus, 263 
Hexapanopeus angustifrons, 111 
Hippolyte 

curacaoensis, 110 

pleuracanthus, 110, 111 

zostericola, 110 
hippos, Caranx, 171, 183 
Hirundichthys speculifer, 172, 184 
hispida, Domecia, 111 
holacanthi, Anilocra, 172, 175, 179 
Holacanthus tricolor, 172, 179 
holocentri, Anilocra, 175, 179 
Holocentrus ascensionis, 172, 179 
hopkinsi, Mesanthura, 47, 51, 267 
Horoloanthura irpex, 267 
horologium, Edwinjoycea, 252, 269 
hummelincki, Cyathura (Stygocyathura), 35 
Hydroniscus quadrifrons, 263 
Hyperphrixus castrensis, 267 
Hypoconcha 

sabulosa, 111 

spinosissima, 111 
Hyporhamphus unifasciatus, 172, 188 
hyptius, Stegophryxus, 113, 268 
Hyssuridae, 16, 58, 60 
Tanirella 

caribbica, 263 

vemae, 263 
Idotea, 255, 259 

balthica, 259 

metallica, 259, 269 
Idoteidae, 251, 252, 255 
Idoteinae, 255, 256 
Iliacantha 

liodactyla, 111 

subglobosa, 111 
imbricata, Parapagurion, 112 
impressa, Politolana, 140 
impressus, Glossobius, 172, 184 
imswe, Kupellonura, 60 
imula, Arubolana, 144, 145 
incisa, Aega, 268 


indica, Limnoria, 194, 195 


infundibulata, Paranthura, 69, 71, 270 


insulae, Limnoria, 195 
insularis 

Alcirona, 158 

Kuna, 170 

Rocinela, 119, 120, 268 
integra, Synsynella, 268 
interger, Cyclograpsus, 111 
intermedius, Palaemonetes, 113 
Ioninae, 107 
Iridopagurus, 112, 113 

iris, 113 
iris, Iridopagurus, 113 
irmae, Haliophasma, 43 
irpex, Horoloanthura, 267 
irrasa, Munida, 111 
irritans, Munidion, 111 
isabelensis, Erichsonella, 269 
Ischnomesidae, 263 
Ischnomesus 

armatus, 263 

caribbicus, 263 

multispinis, 263 
Ischyromene, 210, 214, 217 

barnardi, 218 


itajara, Epinephelus, 122, 172, 190 


jacobus, Myripristis, 172, 180 
jacqueti, Sclerocrangon, 110 
Janiridae, 80, 81, 263 
Janiroidea, 75, 79, 80, 81 
Joeropsidae, 80, 87 
Joeropsis, 87, 88 

bifasciatus, 88 

coralicola, 88, 267 

personatus, 88, 90 

rathbunae, 88, 90, 267, 270 
johanna, Gnathia, 238, 239, 292 
jonesi, Anopsilana, 124, 127 
kadiakensis, Palaemonetes, 113 
kensleyi, Mexicope, 81, 267 
kohleri, Mesosignum, 264 
krebsi, Alcirona, 158, 268, 270 
Kuna, 170, 184 

insularis, 171, 186 
Kupellonura, 60 

formosa, 267 

imswe, 60 
lachneri, Apogon, 171, 188 
Lachnolaimus maximus, 122 
Laminaria, 201 
lamprotaenia, Anchoa, 171, 187 
laodicense, Gnathostenetroides, 78 


lasallae, Astacilla, 252, 253 
lata, Parabopyrella, 112 
lathridia, Amakusanthura, 18, 20 
laticauda, Anilocra, 268 

laticeps, Skuphonura, 58 
latreillei, Tylos, 247, 250 
Latreutes fucorum, 112 


latreuticola, Probopyrinella, 112, 268, 270 


latus, Caranx, 171, 183 
lauffi, Astacilla, 269 
Laurencia, 219 
Leidya 
bimini, 111, 270 
distorta, 111 
Leiostomus xanthurus, 172, 183, 187, 


190 
Lepisosteus spatula, 172, 190 
leptorhynchus, Pandalus, 113 
lewisi, Chalixanthura, 27, 29 
Licranthura, 16, 43 
amyle, 43 
Ligia, 247, 249 
baudiniana, 247, 249 
exotica, 247, 249 
olfersii, 247, 249 
Limnoria, 193, 194, 195, 235 
indica, 194, 195 
insulae, 195 
multipunctata, 195, 196 
pfefferi, 195, 198 
platycauda, 195, 198 
saseboensis, 195, 198 
simulata, 195, 198 
tripunctata, 199 
tuberculata, 194, 195, 199, 268 
unicornis, 195, 199 
Limnoriidae, 114, 193 
lindae, Skuphonura, 267 
linguicauda, Discerceis, 213 
liodactyla, Iliacantha, 111 
Lironeca, 170, 172, 186 
ovalis, 268 
redmanni, 172, 173, 186, 268 
tenuistylis, 171, 186, 187 
texana, 268 
tropicalis, 268 
littoralis 
Eurydice, 148, 149, 268 
Microcerberus, 244 
Loki circumsaltanus, 111 
longicarpus 
Pagurus, 110, 113 
Synalpheus, 110, 111, 113 


INDEX 


longicaudatus, Periclimenes, 113 
longipedis, Munidion, 112, 267 
longipes, Munida, 112 
looensis, Mesanthura, 47, 51 
lunatus, Bothus, 122 
Lutjanus 

analis, 122, 172, 183 

blackfordi, 122 

buccanella, 122 

mahogoni, 172, 183 

synagris, 172, 183 
lyonsi, Edotea, 269 
Lysmata 

rathbunae, 112 

wurdemanni, 112 
Macrobrachium 

acanthurus, 112 

amazonicum, 112 

bonelli, 112 

carcinus, 112 

faustinum, 112 

ohione, 112 

olfersii, 112 

surinamicum, 112 
Macrocystis, 201 
macrostomum, Haemulon, 172, 179 
Macrostylidae, 263 
Macrostylis 

caribbicus, 263 

minutus, 263 

setifer, 264 

vemae, 264 
maculatus 

Scomberomorus, 173, 187 

Sphoeroides, 173, 190 
Madracis, 29, 41, 88, 90, 166 

mirabilis, 29, 41 
magdalenensis, Gnathia, 238, 239 
magnifica, Amakusanthura, 18, 20, 

267 

magnispina, Desmosoma, 263 
mahogoni, Lutjanus, 172, 183 
Malacanthura, 17, 43 

caribbica, 45 

cumanensis, 45 
mangle, Rhizophora, 235 
manningi, Thor, 111 
marcuzzii, Tylos, 247, 250 
margarita, Pontonia, 113 
marginatus, Petrolisthes, 110 
marmoratus, Chiton, 229 
martia, Plesionika, 113 
maximus, Lachnolaimus, 122 


301 


302 INDEX 


mayana 

Bahalana, 128 

Excirolana, 150, 153, 268 
mceclendoni, Synalpheus, 110, 111 
Megalops atlantica, 172, 183 
menziest, Metacirolana, 153, 154 
Mesanthura, 17, 45 

bivittata, 47 

decorata, 53 

fasciata, 47, 49 

floridensis, 53, 267 

hopkinsi, 47, 51, 267 

looensis, 47, 51 

paucidens, 47, 51 

pulchra, 47, 52, 267 

punctillata, 47, 53 

reticulata, 47, 53 
Mesosignidae, 264 
Mesosignum kohleri, 264 
Metacirolana, 144, 153 

agaricicola, 153, 154 

halia, 153, 154 

menziesi, 153, 154 

sphaeromiformis, 153, 154 
metallica, Idotea, 259, 269 
Metaphrixus carolii, 111 
mexicana, Excorallana, 160, 161, 268 
mexicanus, Microcerberus, 269 
Mexicope, 80, 81 

kensleyi, 81, 267 
Mexilana, 273 
mgrayi, Serolis, 202, 268 
Microcerberidae, 243, 244 
Microcerberidea, 14, 243, 269, 273 
Microcerberus, 244, 273 

littoralis, 244 

mexicanus, 269 

minutus, 244 

mirabilis, 244 

nunezi, 244 

renaudi, 244 

simplex, 244 

syrticus, 244 
Microcharon, 90, 91 

herrerai, 93 

phreaticus, 93 

sabulum, 91 
Micropanope barbadensis, 111 
Microparasellidae, 80, 90, 273 
Microphrys bicornutus, 110 
miles, Munida, 112 
millae, Apanthuroides, 27 
milleri, Santia, 99, 267 


minocule, Stenetrium, 100, 102 
minus, Synalpheus, 110, 113 
minuta, Cirolana, 132, 135 
minutus 
Carpias, 82, 84, 270 
Macrostylis, 263 
Microcerberus, 244 
Minyanthura, 16, 53 
corallicola, 53 
mirabilis 
Madracis, 29, 41 
Microcerberus, 244 
Miratidotea, 255, 259 
bruscai, 260 
Monacanthus ciliatus, 172, 190 
montagui, Pandalus, 113 
montosa, Edotea, 269 
morio, Epinephelus, 122 
mortenseni 
Gigantione, 111, 267 
Parabopyrella, 112, 267 


mosaica, Cassidinidea, 208 


motasi, Cyathura (Stygocyathura), 35, 36 


Mothocya, 170, 187 
bohlkeorum, 171, 173, 187, 188 
nana, 172, 187, 188 

Mugil cephalus, 172, 190 

multilineatus, Chromis, 172, 177 

multipunctata, Limnoria, 195, 196 

multispinis, Ischnomesus, 263 

Munida 
constricta, 112 
flinti, 111 
irrasa, 111 
longipes, 112 
miles, 112 
schroederi, 112 
simplex, 110 
stimpsoni, 111 
valida, 110 

Munidion 
cubense, 111 
irritans, 111 
longipedis, 112, 267 

Munna, 93 
petronastes, 94 

Munnidae, 80, 93 

Munnogonium, 96 
wilsoni, 96 

Moycteroperca 
bonaci, 122 
venenosa, 122 

Mpripristis jacobus, 172, 180 


myripristis, Anilocra, 172, 175, 180 
Nalicora, 157, 168 
rapax, 169, 268 
nana, Mothocya, 172, 187, 188 
Nannoniscidae, 264 
Nannoniscus camayae, 264 
narinari, Aetobatus, 166 
Natatolana, 139 
gracilis, 140 
nebulosus, Cynoscion, 172, 183 
Nemanthura, 43 
Neoanthura coeca, 263 
Neopanope 
packardii, 111 
texana sayi, 111 
Neostenetroides, 77, 78 
stocki, 78 
Nerocila, 170, 172, 188 
acuminata, 171, 172, 173, 190, 266, 268 
forma acuminata, 190 
forma aster, 190 
ninae, Armidilloniscus, 246, 247 
niveus, Tylos, 247, 250 
normanni, Alpheus, 112 
northropi, Palaemon, 113 
nunezi, Microcerberus, 244 
nuttingi, Paracerceis, 218, 223 
obtruncata, Cirolana, 132, 135, 268 
occidentalis 
Achelion, 110 
Cleantioides, 256 
Excorallana tricornis, 167 
Parathelges, 112 
ocellatus, Chaetodon, 171, 177 | 
oculata 
Excorallana, 161, 163 
Rocinela, 119, 120, 266, 268 
Oculina arbuscula, 88 
Ocyurus chrysurus, 172, 183 
oestrum, Cymothoa, 171, 172, 173, 182, 183, 
268 
ohione, Macrobrachium, 112 
Olencira praegustator, 268 
olfersii 
Ligia, 247, 249 
Macrobrachium, 112 
Oncilorpheus, 124, 139 
stebbingi, 139 
Oniscidea, 1, 4, 14, 15, 246, 247, 262 
ophiuroides, Echinothambema, 263 
Orbioninae, 107 
orghidani, Cyathura (Stygocyathura), 35, 
36 


INDEX 


Orthopristis 
chrysoptera, 172, 183 
ruber, 122, 173, 179 
ovalis 
Cassidinidea, 207, 208, 268 
Lironeca, 268 
Ovobopyrus alphezemiotes, 267 
oxyophthalmus, Paguristes, 112 
Pachygrapsus transversus, 111 
packardii, Neopanope, 111 
Paguristes 
oxyophthalmus, 112 
tortugae, 112 
Pagurus 
annulipes, 113 
bonairensis, 110, 113 
brevidactylus, 112, 113 
longicarpus, 110, 113 
provenzanoi, 110, 112, 113 
Palaemon 
northropi, 113 
pandaliformis, 113 
Palaemonetes 
exilipes, 113 
intermedius, 113 
kadiakensis, 113 
paludosus, 113 
pugio, 113 
vulgaris, 113 
paludosus, Palaemonetes, 113 
pandalicola, Probopyrus, 112, 266 
pandaliformis, Palaemon, 113 
Pandalus 
annulicornis, 113 
bonnieri, 113 
leptorhynchus, 113 
montagui, 113 
pandionis, Synalpheus, 111, 113 
Panopeus herbstii, 111 
Panoplax depressa, 111 
Parabopyrella 
lata, 112 
mortenseni, 112, 267 
richardsonae, 112, 267 
thomasi, 112 
Parabopyriscus stellatus, 267 
Paracerceis, 210, 218, 223 
caudata, 219, 268, 270 
var. brevipes, 219 
cohenae, 219, 220 
edithae, 218, 221 
glynni, 218, 221 
nuttingt, 218, 223 


303 


304 INDEX 


Paradella, 210, 214, 223 
dianae, 224, 266, 268 
plicatura, 223, 224 
quadripunctata, 223, 224 
tumidicauda, 223, 226 
Paradynamene benjamensis, 268 
Paragnathia, 237 
Paraleptosphaeroma, 207, 208 
glynni, 210, 266 
Paralimnoria, 193, 199 
andrewsi, 199 
forma A, 200 
forma B, 201 
forma typica, 200 
Paraliomera dispar, \11 
Paramunnidae, 80, 96 
Paranthias furcifer, 173, 179 
Paranthura, 64, 69 
antillensis, 69 
barnardi, 69, 71 
floridensis, 69, 71, 267 
infundibulata, 69, 71, 270 
Paranthuridae, 16, 64, 263, 270 
Parapagurion imbricata, 112 
Parapagurus, 112 
parapotamica, Cyathura (Stygocyathura), 35, 36 
Parathelges 
foliatus, 112 
occidentalis, 112 
piriformis, 112, 270 
tumidipes, 112, 270 
partiti, Anilocra, 173, 175, 180 
partitus, Pomacentrus, 173, 180 
parva, Cirolana, 132, 135, 268 
parvioculata, Arubolana, 144, 145 
parviramus, Pseudione, 276 
parvus, Colopisthus, 147, 270 
patulipalma, Stenetrium, 100, 102 
paucidens, Mesanthura, 47, 51 
pectiniger, Synalpheus, 110, 111, 113 
penascoensis, Cortezura, 31 
Pendanthura, 17, 56 
hendleri, 56 
tanaiformis, 56, 270 
penna, Calamus, 122 
perforata, Dynamenella, 213, 215, 270 
Periclimenes 
americanus, 111, 113 
longicaudatus, 113 
periclimenis, Dicropleon, 111 
personata, Eurydice, 147, 149, 270 
personatus, Joeropsis, 88, 90 
petrensis, Eisothistos, 39 


Petrochirus diogenes, 110 
Petrolisthes 

armatus, 110 

galathinus, 110 

marginatus, 110 
petronastes, Munna, 94 
pfefferi, Limnoria, 195, 198 
Phaeoptyx 

conklini, 173, 188 

pigmentaria, 173, 188 
phreaticus, Microcharon, 93 
Phreatocoidea, 14 
Phycolimnoria, 193, 201 

bacescui, 276 

clarkae, 201 
Phyllodurinae, 107 
pigmentaria, Phaeoptyx, 173, 188 
piperata, Eurydice, 147, 149, 268 
piriformis, Parathelges, 112, 270 
planicauda, Cleantioides, 256, 266, 268 
Platybranchiatae, 203 
platycauda, Limnoria, 195, 198 
Plesionika 

antiguai, 113 

edwardsi, 113 

ensis, 113 

heterocarpus, 113 

martia, 113 
pleuracanthus, Hippolyte, 110, 111 
Pleurocope, 97 

floridensis, 98, 267 
Pleurocopidae, 80, 81, 96 
Pleurocrypta floridana, 112 
Pleurocryptella fimbriata, 112 
plicatura, Paradella, 223, 224 
plicifera, Callispongia, 221 
plumieri, Haemulon, 172, 179 
Pogonias cromis, 173, 190 
polita 

Cyathura, 267 

Politolana, 140, 143, 268 
Politolana, 139, 140 

impressa, 140 

polita, 140, 143, 268 
Pomacentrus partitus, 173, 180 
Pontonia margarita, 113 
Porcellana sayana, 110 
Porites, 88, 90 
poteriophora, Akidognathia, 264 
praegustator, Olencira, 268 
Priacanthus arenatus, 173, 183 
probatocephalus, Archosargus, 122 
Probopyria alphei, 112, 267 


Probopyrinella 

heardi, 267 

latreuticola, 112, 268, 270 
Probopyrus pandalicola, 112, 266 
Processa 

acutirostris, 113 

canaliculata, 113 

edulis, 113 

JSimbriata, 113 

tenuipes, 113 
processae, Urobopyrus, 113, 268 
Prodajus cf. bigelowiensis, 268 
productatelson, Exosphaeroma, 229, 231 
provenzanoi, Pagurus, 110, 112, 113 
psamathus, Angliera, 91 
Pseudasymmetrione, 113 
Pseudione 

affinis, 113, 270 

cognata, 268 

parviramus, 276 

upogebiae, 268 
Pseudioninae, 107 
Ptilanthura tricarina, 267 
puertoricensis, Gnathia, 238, 241 
pugilator, Uca, 111 
pugio 

Gnathostenetroides, 77, 267 

Palaemonetes, 113 
pulchra 

caribbea, Storthyngura, 263 

Mesanthura, 47, 52, 267 
punctatus, Carpias, 82, 85 
punctillata, Mesanthura, 47, 53 
purpureus, Arcturus, 264 
Pylopagurus, 110 

corallinus, 112 
quadricornis, Excorallana, 161, 165, 270 
quadridentata, Sphaeroma, 234, 268 
quadrifrons, Hydroniscus, 263 
quadrilirata, Dynamenella, 213, 215 
quadripunctata, Paradella, 223, 223 
racovitzai, Angliera, 91 
radicicola 

Anopsilana, 124, 127 

Xylolana, 157 
Raja eglanteria, 122 
rapax, Nalicora, 169, 268 
rathbunae 

Joeropsis, 88, 90, 267, 270 

Lysmata, 112 
rathi, Gnathia, 238, 241 
redmanni, Lironeca, 172, 173, 186, 268 
regalis, Scomberomorus, 173, 187 


INDEX 


regina, Astacilla, 252, 253 
renaudi, Microcerberus, 244 
Renocila, 170, 191 

bowmani, 173, 191 

colini, 171, 191 

waldneri, 173, 191, 193 
reticulata, Mesanthura, 47, 53 
reynoldsi, Uromunna, 94, 95, 266, 267 
Rhamphion, see Aega (Ramphion) 
Rhizophora mangle, 235 
rhombeus, Gerres, 172, 187 
Rhyscotus, 247, 249 

texensis, 247, 249 
richardsonae, Parabopyrella, 112, 267 
ricordi, Sesarma, 111 
riedli, Vandeloscia, 247, 251 
Rithropanopeus harrisii, 111 
Rocinela, 116, 119 

cubensis, 119, 120 

insularis, 119, 120, 268 

oculata, 119, 120, 266, 268 

signata, 119, 120, 266, 268 
rostrata, Galathea, 112 
ruber 

Caranx, 171, 183 

Orthopristis, 122, 173, 179 
ruetzleri, Cymodoce, 227 
sabulosa, Hypoconcha, 111 
sabulum, Microcharon, 91 


305 


salpiscinalis, Cyathura (Stygocyathura), 35, 38 


Samariensis 

Edotea, 276 

Gnathia, 238, 251 
Santia, 98 

milleri, 99, 267 
Santiidae, 80, 98 
Sargassum, 84, 201 
saseboensis, Limnoria, 195, 198 
Saxatilis, Abudefduf, 171, 175, 186 
sayana, Porcellana, 110 
sayi, Neopanope texana, 111 
sbordonii, Cyathura (Stygocyathura), 35, 

38 

Schizobopyrina urocaridis, 113, 268 
schmitti, Azygopleon, 110, 267 
schoepfi 

Alutera, 171, 190 

Chilomycterus, 172, 190 
schroederi, Munida, 112 
sciurus, Haemulon, 172, 179 
Sclerocrangon jacqueti, 110 
Scomberomorus 

cavalla, 173, 187 


306 INDEX 


Scomberomorus (cont.) 

maculatus, 173, 187 

regalis, 173, 187 
scopulosa, Chalixanthura, 27, 29 
sedentarius, Chaetodon, 171, 177 
Selar crumenophthalmus, 173, 183 
Serolidae, 114, 115, 201 
Serolis, 202 

mgrayi, 202, 268 
Serranus tigrinus, 173, 191, 193 
serrata, Spinianirella, 263 
serratum, Stenetrium, 100, 102 
serricaudus, Carpias, 82, 87 
Sesarma ricordi, 111 
seticornis, Stenorhynchus, 110 
setifer, Macrostylis, 264 
Setosa, Accalathura, 64, 65 
sexticornis, Excorallana, 161, 165 
signata 

Amakusanthura, 18, 21 

Rocinela, 119, 120, 266, 268 
significa, Amakusanthura, 18, 23 
simplex 

Microcerberus, 244 

Munida, 110 
simulata, Limnoria, 195, 198 
Skuphonura, 17, 58 

laticeps, 58 

lindae, 267 
snanoi, Storthyngura, 263 
somala, Haptolana, 138 
Sparisoma viride, 122 


spathulicarpus, Stenetrium, 100, 104 


Spatula, Lepisosteus, 172, 190 
speculifer, Hirundichthys, 172, 184 


specus, Cyathura (Stygocyathura), 35, 38 


Speocirolana, 273 

Sphaerolana, 273 

Sphaeroma, 226, 232, 234 
annandalei, 234 
destructor, 235 
quadridentata, 234, 268 
terebrans, 234, 235, 268 
walkeri, 234, 235 


Sphaeromatidae, 114, 115, 202, 204 


Sphaeromatinae, 204, 226 

sphaeromiformis, Metacirolana, 153, 
154 

Sphoeroides maculatus, 173, 190 

Sphyraena barracuda, 122 

Spinata, Arcturella, 252, 269 

Spinianirella serrata, 263 


spinosissima 

Acanthocope, 263 

Hypoconcha, 111 
stebbingi 

Oncilorpheus, 139 

Stenetrium, 100, 104, 267, 270 
Stegias clibanarii, 113, 270 
Stegophryxus hyptius, 113, 268 
steindachneri, Haemulon, 122 
stellatus 

Astrapogon, 171, 188 

Parabopyriscus, 267 
Stenetriidae, 99 
Stenetrioidea, 99 
Stenetrium, 99, 100 

bowmani, 100 

minocule, 100, 102 

patulipalma, 100, 102 

serratum, 100, 102 

Spathulicarpus, 100, 104 

stebbingi, 100, 104, 267, 270 
Stenobermuda, 99, 106 

acutirostrata, 106, 270 
Stenorhynchus seticornis, 110 
stenotelson, Anthomuda, 23, 270 
Stimpsoni, Munida, 111 
stocki, Neostenetroides, 78 
Storthyngura 

pulchra caribbea, 263 

snanoi, 263 
Striata, Yvesia, 246 
striatus, Chaetodon, 171, 177 
Stygocyathura, see Cyathura (Stygocyathura) 
subcaudalis, Eophrixus, 111, 267 
subglobosa, Iliacantha, 111 
subtilis, Excorallana, 160 
sulcipes, Dactylokepon, 267 
surinamensis, Batrachoides, 171, 190 
surinamicum, Macrobrachium, 112 
symmetricus, Ambidexter, 113 
synagris, Lutjanus, 172, 183 
synalphei 

Bopyrione, 110, 267 

Hemiarthrus, 111, 267 
Synalpheion giardi, 113 
Synalpheus 

bousfieldi, 110 

brevicarpus, 110 

brooksi, 110, 111, 113 

Sritzmuelleri, 110, 111 

goodei, 110, 111 

hemphilli, 110, 111 


longicarpus, 110, 111, 113 
meclendoni, 110, 111 
minus, 110, 113 
pandionis, 111, 113 
pectiniger, 110, 111, 113 
Synodus foetens, 173, 183 
Synsynella, 110, 113 
choprae, 113, 268, 270 
deformans, 110, 113, 268, 270 
integra, 268 
Syringodium, 166, 232 
filiforme, 253, 260 
syrticus, Microcerberus, 244 
tanaiformis, Pendanthura, 56, 270 
Tecticeps, 204 
Tecticipitinae, 204 
tenuipes 
Aega, 117, 119 
Processa, 113 
tenuis, Colanthura, 65, 270 
tenuistylis, Lironeca, 171, 186, 187 
terebrans, Sphaeroma, 234, 235, 268 
teri, Eisothistos, 39 
testudinea, Thalassia, 251 
texana, Lironeca, 268 
texensis, Rhyscotus, 247, 249 
Thalassia, 166, 232 
testudinea, 251 
Thermarcturus, 252, 255 
venezuelensis, 255 
thomasi, Parabopyrella, 112 
Thor 
floridanus, 110, 111 
manningi, 111 
thorii, Bopyrinella, 110 
tigrinus, Serranus, 173, 191, 193 
tortugae, Paguristes, 112 
tortuganus, Balanopleon, 110 
Tozeuma carolinense, 112 
transversa, Ceratothoa, 268 
transversus, Pachygrapsus, 111 
tricarina, Ptilanthura, 267 
tricarinata, Booralana, 275 
trichostoma, Haptolana, 138 
tricolor 
Clibanarius, 110, 112, 113 
Holacanthus, 172, 179 
tricornis, Excorallana, 161, 165, 266, 268 
occidentalis, 167 
tricornis, 161, 165 
Tridentella, 236 
virginiana, 236 


INDEX 


Tridentellidae, 115, 235, 236 
triospathiona, Gnathia, 238, 241 
tripunctata, Limnoria, 199 
triton, Carpias, 82, 87 
Troglocirolana, 273 
tropicalis 

Erichsonella filiformis, 257 

Haplomesus, 263 

Lironeca, 268 
tuberculata, Limnoria, 194, 195, 199, 268 
tuberculatus, Chiton, 229 
tumidicauda, Paradella, 223, 226 
tumidipes, Parathelges, 112, 270 
Turbinaria, 166, 219 
Tylos, 247, 250 

latreillei, 247, 250 

marcuzzit, 247, 250 

niveus, 247, 250 

wegeneri, 247, 250 
typica, var., Dynamenella acutitelson, 214 
uberlackerae, Gigantione, 267 
Uca, 111 

pugilator, 111 
unicornis 

Haploniscus, 263 

Limnoria, 195, 199 
unifasciatus, Hyporhamphus, 172, 188 
univam, Cyathura (Stygocyathura), 35, 

38 

Upogebia affinis, 112 
upogebiae, Pseudione, 268 
Urobopyrus processae, 113, 268 
urocaridis, Schizobopyrina, 113, 268 
Uromunna, 93, 94 

caribea, 94 

hayesi, 267 

reynoldsi, 94, 95, 266, 267 
valeriae, Haliophasma, 41, 42 
valida, Munida, 110 
Valvifera, 14, 251, 264, 269 
Vandeloscia, 247, 251 

culebrae, 247, 251 

riedli, 247, 251 
vanderhorsti, Clastotoechus, 110 
velosa, Gnathia, 238, 243 
vemae 

Tanirella, 263 

Macrostylis, 264 
venenosa, Mycteroperca, 122 
Venezanthura, 31 
venezuelensis, Thermarcturus, 255 
vetula, Balistes, 122 


307 


308 INDEX 


Virganthura, 64, 73 

crassa, 73 
virginalis, Gnathia, 238, 243 
virginiana, Tridentella, 236 
viridari, Alpheus, 112 
viride, Sparisoma, 122 
vittatus, Clibanarius, 110, 112 
vulgaris, Palaemonetes, 113 
waldneri, Renocila, 173, 191, 193 
walkeri, Sphaeroma, 234, 235 
warmingii, Excorallana, 161, 167 
wegeneri, Tylos, 247, 250 
wilsoni, Munnogonium, 96 


wolffi, Bopyrissa, 110 
wurdemanni, Lysmata, 112 
xanthurus, Leiostomus, 172, 183, 187, 190 
Xenanthura, 60 
brevitelson, 62, 267 
Xylolana, 123, 143, 144, 156 
radicicola, 157 
yucatanensis, Bythognathia, 275 
yucatanum, Exosphaeroma, 229, 231 
Yuesia, 246 
Striata, 246 
zostericola, Hippolyte, 110 


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